Biology
590.5 Fl N.S.
no. 104(2005)
AUG ^ 3 2005
HI
^[yr\
Zoology
NEW SERIES, NO. 104
Systematic Review of Japanese Macaques,
Macaca fuscata (Gray, 1870)
Jack Fooden
Mitsuru Aimi
March 14, 2005
Publication 1533
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UNIVERSITY OF ILLINOIS
LIBRARY
AT URBANA-CHAMPAIGN
BIOLOGY
FIELDIANA
Zoology
NEW SERIES, NO. 104
Systematic Review of Japanese Macaques,
Macaca fuscata (Gray, 1870)
Jack Fooden
Mitsuru Aimi
Division of Mammals
Department of Evolution and Phylogeny
Department of Zoology
Primate Research Institute
Field Museum of Natural History
Kyoto University
1400 South Lake Shore Drive
Inuyama, Aichi 484-8506
Chicago, Illinois 60605-2496
Japan
U.S.A.
Accepted August 3, 2004
Published March 14, 2005
Publication 1533
BIOLOGY LIBRARY
101 BURRILL HALL
AUG 0 3 2005
PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY
© 2005 Field Museum of Natural History
ISSN 0015-0754
PRINTED IN THE UNITED STATES OF AMERICA
Table of Contents
Abstract 1
Introduction 1
Geographic Distribution, Total Popula-
tion Estimate 3
External Characters 5
Pelage 5
General Characterization 5
Early Development 5
Seasonal Variation 5
Geographic Variation 6
Albinism 8
External Measurements and Proportions .. 8
Linear Measurements of Body Size .... 8
Tail Length 11
Hind Foot Length 12
Body Weight 13
Summary 13
Dermatoglyphics 15
Congenital Malformation of Limbs 15
Cranial Characters 17
Effect of Provisioning 17
Sex and Age Variation 17
Geographic Variation 19
Interspecific Comparisons 21
Molecular Biology and Genetics 22
DNA: Intraspecific Studies 22
Mitochondrial DNA 22
Minisatellite DNA and Microsatellite
DNA 33
DNA: Interspecific Studies 33
Mitochondrial DNA 33
Nuclear DNA: Noncoding Intron 33
Nuclear DNA: Y-Chromosome 35
Nuclear DNA: Random Amplified Poly-
morphism 35
Mitochondrial DNA-Nuclear DNA:
Multiple Loci 35
Blood Proteins 37
Hemoglobin Amino- Acid Sequence 42
Antibodies to Cedar Pollen Antigens 43
Karyology 45
Parasites 46
Protistans 46
Trematodes 47
Cestodes 47
Nematodes 47
Mite 49
Lice 49
Tick 49
Q Fever Bacterium 49
Viruses 49
Natural History 50
Habitats 50
Terrestriality/Arboreality 51
Group Size and Composition 51
Home Range Area and Group Density ... 52
Diet 52
Predators 54
Intergroup Dispersal 54
Reproduction 54
Seasonality 54
Sexual Maturation 57
Sexual Skin 57
Menstrual Cycle 58
Estrus 58
Consortship 59
Copulatory Behavior 60
Dominance Rank and Reproductive Suc-
cess 60
Inbreeding 61
Hybridization 61
Nonreproductive Sexual Behavior 62
Gestation 63
Parturition 64
Neonatal Sex Ratio; Birth Weight 65
Twinning 66
Birthrate 66
Infant Mortality 68
Nursing; Weaning 68
Longevity; Reproductive Senescence 71
Population Growth Rate 72
Fossils 72
S ystematics 72
Geographic Variation and Subspecific
Recognition 72
Pelage Color 75
Body Size 75
Cranial Morphology 75
Mitochondrial DNA; Blood Proteins ... 75
Key to External Characters of Recog-
nized Subspecies 77
Japanese Vernacular Names Applied to
Macaca fuscata 77
Subspecies Accounts:
Macaca fuscata fuscata (Gray, 1870) 77
Synonymy 77
Type Series 79
Type Locality 80
Distribution 81
Diagnosis 81
Remarks 81
Specimens Examined 82
Macaca fuscata yakui Kuroda, 1 940 82
Synonymy 82
Type Series 82
ui
Type Locality 83
Distribution 83
Diagnosis 83
Specimens Examined 83
Evolution and Dispersal 83
Acknowledgments 86
Literature Cited 86
Appendix L Specimens Examined 105
Appendix 2: Macaca fuscata Localities
Reported in Postal Survey' Conduct-
ed BY K. Hasebe in 1923 108
Appendix 3. Locality Records (Specimens
Examined and Literature Records)
OF Macaca fuscata 113
Appendix 4: Gazetteer of Macaca fuscata
Collection and Observation Locali-
ties 119
Appendix 5: Local Variation in Sitting
Height in Macaca fuscata Adults on
Honshu 191
Appendix 6: Local Variation in Anterior
Trunk Length in Macaca fuscata
Adults on Honshu 192
Appendix 7: Local Variation in Tail
Length and Tail Length/Anterior
Trunk Length ratio in Macaca fusca-
ta Adults on Honshu 194
Appendix 8: Local Variation in Hind
Foot Length and Hind Foot/Anteri-
or Trunk Length Ratio in Macaca
fuscata Adults on Honshu 196
Appendix 9: Local Variation in Body
Weight in Macaca fuscata Adults on
Honshu 197
Appendix 10: Local Variation in Great-
est Length of Skull in Macaca fus-
cata Adults on Honshu 198
Index 199
Note Added in Proof 200
List of Illustrations
1. Macaca fuscata depicted in first scroll
(detail) of four-scroll Choju-jinbutsu-
giga set 2
2. Survey reports of geographic distribu-
tion of Macaca fuscata
A. Survey conducted in 1923 by K.
Hasebe 4
B. Survey conducted in 1992 by Envi-
ronmental Agency of Japan 4
3. Macaca fuscata — adult male, adult fe-
male, and infant — photographed at
Takasakiyama, Oita Prefecture,
Kyushu 6
4. Latitudinal variation in head and body
length in Honshu and Shikoku samples
of Macaca fuscata aduhs, 10
5. Latitudinal variation in sitting height in
Macaca fuscata adults 12
6. Latitudinal variation in anterior trunk
length in Macaca fuscata adults 14
7. Latitudinal variation in tail length in
Macaca fuscata adults 16
8. Tail length vs. head and body length in
Macaca fuscata immatures and adults
' 18
9. Latitudinal variation in hind foot length
m Macaca fuscata aduXis 20
10. Latitudinal variation in body weight in
Macaca fuscata adults 22
1 1 . Skull of adult female Macaca fuscata
fuscata 26
12. Skull of adult female Macaca fuscata
yakui 27
13. Skull of adult male Macaca fuscata
fuscata 28
14. Skull of adult male Macaca fuscata
yakui 29
15. Latitudinal variation in greatest length
of skull in Macaca fuscata adults 32
16. Greatest length of skull vs. estimated
age in Honshu and Yakushima speci-
mens of Macaca fuscata 34
17. Orbital height vs. orbital breadth in in-
sular samples of Macaca fuscata adults
36
18. Geographic variation in mtDNA se-
quence haplotypes in Macaca fuscata
(Hayasaka et al., 1991) 38
19. Geographic variation in mtDNA se-
quence haplotypes in Macaca fuscata
(Kawamoto, 2002) 41
20. Geographic distribution and genetic in-
terrelationships of blood-protein groups
in Macaca fuscata 43
21. Distribution oi Macaca fuscata troops
in Japan for which birth seasonality
data are available 55
22. Latitudinal variation in birth seasonali-
ty in in-situ troops of Macaca fuscata
56
23. Age-specific birthrates in two nonpro-
visioned troops and two provisioned
troops of Macaca fuscata 70
IV
24. Distribution of fossil macaque locali-
ties in Japan 76
25. Bathymetric map of Yakushima-Sakha-
lin island-chain area, illustrating proba-
ble land bridges exposed by sea-level
depression of 140 m 84
List of Tables
1 . Estimates of total extant population of
Macaca fuscata 5
2. Geographic variation in mean paleness
index values of pelage on dorsal sur-
face of trunk, dorsal surface of hands,
and dorsal surface of feet in Macaca
fuscata 7
3. Months during which molting was ob-
served in nine Macaca fuscata captives
maintained outdoors at the Japan Mon-
key Centre, Inuyama, Aichi Prefecture,
Honshu 7
4. Dorsal hair density in infant and adult
Macaca fuscata 8
5. Prefectural variation in head and body
length and related external measure-
ments in Macaca fuscata 9
6. Insular variation in sitting height in
Macaca fuscata adults 11
7. Insular variation in anterior trunk
length in Macaca fuscata adults 13
8. Insular variation in tail length and tail
length/anterior trunk length ratio in
Macaca fuscata adults 15
9. External measurements in seven Maca-
ca fuscata adults with unusually short
tails 17
10. Insular variation in hind foot length
and hind foot length/anterior trunk
length ratio in Macaca fuscata adults .. 19
1 1 . Insular variation in body weight in
Macaca fuscata adults 21
12. Frequency of dermatoglyphic patterns
on both hands in five local samples of
Macaca fuscata 23
13. Provisioning status and occurrence of
congenital limb malformations in 65
Macaca fuscata groups 23
14. Neonatal incidence of congenital limb
malformations in nine Macaca fuscata
groups 24
15. Sexual variation in incidence of con-
genital limb malformations in Macaca
fuscata neonates 24
16. Greatest length of skull in sample of
provisioned Macaca fuscata population
collected at Takagoyama, Chiba Prefec-
ture, Honshu, compared with that in
sample of nonprovisioned populations
collected elsewhere in Chiba Prefecture
30
17. Cranial measurements and proportions
in age/sex classes of nonprovisioned
wild-collected Macaca /Micara 30
18. Dental emergence chronology in Maca-
ca fuscata 31
19. Insular variation in greatest length of
skull (mm) in wild-collected Macaca
fuscata 31
20. Insular variation in head length (great-
est midsagittal length from glabella to
occiput) in living Macaca fuscata 33
21. Insular variation in orbital measure-
ments and orbital height/breadth index
in adult wild-collected Macaca fuscata
35
22. Greatest length of skull and rostral-
postrostral ratio \n fascicularis-gxoxyp
species 37
23. Local variation in mtDNA sequence
haplotypes in Macaca fuscata (Hayasa-
kaet al., 1991) 39
24. Mitochondrial DNA haplotypes detect-
ed in Macaca fuscata at localities sam-
pled by Kawamoto (2002) 40
25. Mean mtDNA divergence estimates re-
ported between Macaca fuscata and M.
mulatta, M. cyclopis, and M. fascicu-
laris 42
26. Sister-group relationships of Macaca
fuscata indicated by sequencing of pa-
ternal, maternal, and biparental markers
42
27. Mean frequency of major alleles at 2
loci in blood-protein groups of Macaca
fuscata and frequency of corresponding
alleles in southern Chinese M. mulatta
44
28. Incidence of serum specimens positive
for antibodies to crude cedar pollen an-
tigen and incidence of serum speci-
mens with positive reactions to purified
cedar pollen antigens in prefectural
samples of Macaca fuscata 46
29. Incidence of protistan parasite species
detected in fecal samples of Macaca
fuscata on Shimokita Peninsula, Ao-
mori Prefecture 46
30. Incidence of cestode parasites in local
samples of Macaco fuscata 47
3 1 . Incidence of nematode parasites in lo-
cal samples of Macaco fiiscata 48
32. Incidence of antibodies to eight viruses
in three local groups of Macaco fusca-
ta ^ 50
33. Relationship between housing system
and incidence of antibodies to hepatitis
A in captive Macaco fiiscata 51
34. Geographic variation in group size in
wild Mococa fuscata populations 52
35. Group composition and ratios in wild
Macaco fuscata 53
36. Menstrual cycle length in Macaco fus-
cata 58
37. Frequency and duration of estrous peri-
ods in mature Macaco fuscata females
59
38. Reported results concerning relation-
ship between male dominance rank and
reproductive success in natural and
seminatural groups of Macaco fuscato
60
39. References concerning nonreproductive
sexual behavior reported in natural and
seminatural groups of Macaco fuscata
62
40. Gestation period duration in Macaco
fuscata 64
41. Neonatal sex ratio in Macaco fuscata .. 65
42. Birth weight in Afacaca yzY5cara 66
43. Annual birthrate in Macaca yM5cara .... 67
44. Effect of Macaco fuscata infant death
on interbirth interval of mother 68
45. Relationship between birthrate and fe-
male dominance rank in nonprovi-
sioned and provisoned groups of Ma-
coca fuscato 68
46. Infant mortality rate in nonprovisioned
and provisioned groups of Macaco fus-
cato 69
47. Mean life span in Macaco fuscato fe-
males 71
48. Population growth rate in Mococa fiis-
cata groups 73
49. Fossil macaque localities in Japan,
with indication of associated fossil pro-
boscideans 74
50. Mammal stages, biozones. and stratig-
raphy of Quaternary deposits in Japan
75
51. Dental measurements in four Korean
fossil macaque specimens collected at
Turubong Cave and one Japanese fossil
macaque specimen collected at Ando
Quarry 77
52. Dental measurements in samples of ex-
tant Mococa fuscata, for comparison
with corresponding measurements in
Korean and Japanese fossil macaques 78
53. Distribution relative to Tsugaru Strait
(between Honshu and Hokkaido) of
Yakushima-Sakhalin species of nonvo-
lant terrestrial mammals 83
VI
Systematic Review of Japanese Macaques,
Macaca fuscata (Gray, 1870)
Jack Fooden
Mitsuru Aimi
Abstract
Japanese macaques, Macaca fuscata (Gray, 1 870), are systematically reviewed, based on
examination of 1264 specimens, survey of relevant literature, and observation of natural pop-
ulations. This review includes analyses of external and cranial characters, molecular biology
and genetics, and parasites. Information also is presented concerning natural history, reproduc-
tion, fossils, and taxonomic history. Concerning the evolutionary history of M. fuscata, avail-
able evidence suggests that this species is derived from a M. mulatta-like population that
dispersed to the Japanese archipelago during one or two glacial intervals of sea-level depres-
sion, ca. 0.63-0.43 million years ago (Ma), via a now-submerged dry-land connection between
the Korean Peninsula and the Kyushu-West Honshu area. In an appendix, an annotated gaz-
etteer lists 1347 localities at which M. fuscata has been collected or observed.
Introduction
Macaca fuscata is a stump-tailed monkey that
is unique to the Japanese archipelago and is the
only nonhuman primate that inhabits these is-
lands. Its geographic distribution, which extends
from ca. 30°15'N to 4r30'N, is the northernmost
of any extant nonhuman primate.
Although Japanese artists have been fascinated
by the morphology and behavior of M. fuscata for
at least eight centuries (Fig. 1; cf. Ohnuki-Tier-
ney, 1987, p. 27), Western science apparently did
not become aware of the existence of this species
until 1727, when it was briefly mentioned by E.
Kaempfer (1727, p. 126; cf. Bodart-Bailey, 1999,
p. 72) in his book The History of Japan . . . ,
Kaempfer, a German physician, encountered M.
fuscata while serving with a Dutch trading mis-
sion in Japan during the period 1690-1692. The
first scientific specimens of M. fuscata were sent
to Europe in 1821-1822 by C. G. Reinwardt (let-
ters, RMNH archives) and in 1823-1829 by R E
von Siebold (Holthuis & Sakai, 1970, p. 25). The
first scientific account of the species was pub-
lished in 1825 by E Cuvier (1824-1829, livraison
47, p. 1), based on a captive exhibited in a zoo in
India.
The present review of M. fuscata is based on
study of 1264 specimens (Appendix 1), survey of
relevant literature, and observation of natural pop-
ulations. Specimens examined are preserved in in-
stitutions listed below, which hereafter are cited
by means of the indicated abbreviations; the num-
ber of M. fuscata specimens studied at each in-
stitution is indicated by a parenthetical notation.
AIUM Anthropologisches Institut der Univer-
sitat Miinchen (32)
AIUZ Anthropologisches Institut der Univer-
sitat Zurich (2)
AMNH American Museum of Natural History,
New York (3)
ANSP Academy of Natural Sciences, Phila-
delphia (3)
BM(NH) The Natural History Museum, London
(12)
JMC Japan Monkey Centre, Inuyama, Japan
(39)
KFL Koshima Eield Laboratory, Kushima,
Japan (48)
MCZ Museum of Comparative Zoology, Har-
vard University, Cambridge, Massachu-
setts (2)
FIELDIANA: ZOOLOGY, N.S., NO. 104, MARCH 14, 2005, PP. 1-200
f
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Fig. 1. Macaca fuscata depicted in first scroll (detail) of four-scroll Choju-jinbutsu-giga set, in collection of Kozanji
monastery. Kyoto (cf. Ohnuki-Tierney, 1987, p. 28); attributed to Toba Sojo, also known as Kakuyu (1053-1140),
noted priest and artist. (Reproduced by permission of Kozanji monastery.)
FIELDIANA: ZOOLOGY
MNHN Museum National d'Histoire Naturelle,
Paris (5)
MUE Miyagi University of Education, Sen-
dai, Japan (15)
NHR Naturhistoriska Riksmuseet, Stockholm
(2)
PRIKU Primate Research Institute, Kyoto Uni-
versity, Inuyama, Japan (895)
RMNH Nationaal Naturhistorisch Museum,
Leiden (14)
SU Saitama University, Urawa, Japan (96)
TPM Tochigi Prefectural Museum, Utsuno-
miya, Japan (32)
USNM National Museum of Natural History,
Washington, D.C. (12)
ZMB Museum fiir Naturkunde, Berlin (28)
ZMUZ Zoologisches Museum der Universitat
Zurich (2)
ZSBS Zoologisches Staatssammlung, Munich
(22)
Geographic Distribution, Total
Population Estimate
The most recent comprehensive survey of the
geographic distribution of M. fuscata was con-
ducted in 1992 by the Environmental Agency of
Japan (now Ministry of the Environment, Japan).
In this survey, the land area of the Japanese ar-
chipelago was subdivided into a grid of 10 X 10-
km cells, for each of which the presence or ab-
sence of monkeys was reported by local officials
(Fig. 2B); M. fuscata was reported present in ca.
1000 of the 10 X 10-km cells. This survey con-
firms that M. fuscata is relatively widely distrib-
uted in upland areas on three of the four large
Japanese islands — Honshu, Shikoku, and Kyu-
shu— and that it also inhabits six close-lying small
islands — Kinkazan (off northeastern Honshu),
Awajishima and Shodoshima (in the Inland Sea,
between Honshu and Shikoku), Kashima (off
southwestern Shikoku), Kojima {- Koshima, off
southeastern Kyushu), and Yakushima (ca. 60 km
south of Kyushu); all six of the small islands in-
habited by M. fuscata are within the 100-m bathy-
metric line (Iwano, 1974, figs. 1-5). M. fuscata
formerly inhabited three additional small islands
where it is now extinct (Appendix 3; Kawanaka,
1973, p. 115); these three islands are Omishima
(north of westernmost Honshu), where M. fuscata
survived until ca. 1965 (Tanaka, 1991, p. 19); Hi-
radoshima (northwest of Kyushu), where this spe-
cies survived until ca. 1925 (Sakura, 1976, p.
151); and Tanegashima (northeast of Yakushima),
where it survived until ca. 1955 (Azuma, 1972, p.
261) and subsequently became extinct as a result
of deforestation.
The earliest comprehensive survey of the dis-
tribution of M. fuscata was conducted in 1 923 by
K. Hasebe, but the results of this survey were not
published until 1974 (Iwano, 1974, p. 5; cf. Ama-
gasa & Ito, 1978, p. 96); somewhat less rigorous
surveys were conducted by K. Kishida in 1953
and by H. Takeshita in 1964a,b (Iwano, 1974, p.
8). Hasebe mailed 563 survey questionnaires to
heads of each prefecture and county in Japan and
received 535 replies that identified a total of 818
monkey localities (Fig. 2A). Although Hasebe's
survey preceded the Environmental Agency sur-
vey (Fig. 2B) by nearly seven decades, the two
surveys reveal basically similar patterns of geo-
graphic distribution; in particular, both surveys in-
dicate relatively few records of monkeys in three
areas that are within the geographic range of M.
fuscata — northern Honshu (ca. 39-40. 5°N), north
of Boso Peninsula (ca. 35.5-36.5°N), and north-
western Kyushu (ca. >33°N, <131°E). A map of
localities of M. fuscata specimens and literature
records examined for the present study (Appendix
3; cf. Baldwin et al., 1980, p. 269) also reveals a
pattern of geographic distribution generally simi-
lar to that revealed by the 1 992 survey map. Fac-
tors responsible for the sparse distribution of M.
fuscata in northern Honshu (ca. 39-40. 5°N) have
been analyzed historically by Mito (1992, p. 143;
cf. Oi et al., 1997, p. 9; 2000, p. 11).
Although M. fuscata is now absent in most
lowland areas of Japan (Amagasa & Ito, 1978, p.
102), it presumably inhabited lowland areas be-
fore their intensive exploitation by humans (cf.
Tokita et al., 1978, p. 88). The greatest elevation
at which M. fuscata has been reported is 3 1 80 m
(see below. Natural History: Habitats).
M. fuscata is absent on Hokkaido (the north-
ernmost of the four large Japanese islands), on the
Ryukyu archipelago (southwest of Kyushu and
Yakushima), and on most small islands that sur-
round Honshu, Shikoku, and Kyushu — including
most islands within the 100-m bathymetric line
(Iwano, 1974, figs. 1-5). One small island — Mi-
yajima — is inhabited by an artificially introduced
population of M. fuscata, and two other small is-
lands— Okinoshima, and Tomogashima — former-
ly were inhabited by introduced populations (for
details, see Gazetteer, Appendix 4). Fragmentary
skeletal evidence indicates that M. fuscata was
FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES
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HELDIANA: ZOOLOGY
Table 1 . Estimates of total extant population of Macaca fuscata.
Estimated
Estimated
total
number of
References
population
groups
Basis of estimates
Kishida(1953, p. 65)
15,614
313
Questionnaire survey, 1950
Takeshita (1964b, p. 20)
22,000-34,000
425
Questionnaire survey, 1961-1962
Kawamura (1973, p. 456)
43.161
—
Questionnaire survey,' 1970
Kawamura (1975, p. 502)^
50,000-100,000
—
Field survey (incomplete)
Hashiba(1989, p. 30)
114,431
—
Prefectural survey of broadleaf
forest areas
Survey conducted by H. Takeshita.
Cf. Kawamura (1973, p. 455).
translocated from Yakushima to Okinawa (Shuri
Castle) several centuries ago (Mouri et al., 2000,
p. 87); artificially modified monkey bones discov-
ered at an archaeological site on Rebun Island,
northwest of Hokkaido, may have been transport-
ed to Rebun Island as artifacts (Nishimoto, 1981,
p. 430).
Prior to ca. 1945, the geographic distribution of
M. fuscata apparently was contracting (K. Watan-
abe and Y. Muroyama, PRIKU, pers. comm.).
Subsequently— despite recent annual extermina-
tion of ca. 10,000 monkeys as agricultural pests—
the distribution generally has been expanding; this
contraction and expansion apparently are related
to changing patterns of Japanese human demog-
raphy and behavior (cf. Sprague, 2002, p. 255).
The extant total population of M. fuscata is esti-
mated to be approximately 100,000 individuals
(Table 1).
External Characters
Pelage (Fig. 3)
General Characterization — In M. fuscata
adults in prime pelage, the color of the crown and
dorsal surface of the trunk varies from pale yel-
lowish brown to grayish brown to dark golden
brown. Fur on the outer surface of the limbs and
tail tends to be slightly paler than that on the ad-
jacent surface of the trunk (cf. Hamada et al.,
1992, p. 6); fur on the dorsal surface of the hands
varies from pale yellowish brown to blackish, and
that on the feet varies from pale yellowish brown
to dark brown. Fur on the ventral surface of the
trunk and the inner surface of the limbs varies
from whitish to pale gray to pale ochraceous buff.
The prominent side-whiskers are somewhat paler
than the crown, and the lateral facial crest pattern
is infrazygomatic (Fooden, 1995, p. 18). The thin-
ly haired facial skin is pinkish to reddish (for dis-
cussion of sexual skin, see below. Reproduction).
Dorsal pelage color in adult females averages
slightly paler than in adult males (Table 2; Ha-
mada et al., 1992, p. 5). In a very old free-ranging
male (estimated age 38 years), the fur on the
haunches and dorsal surface of the trunk report-
edly had become "quite white" (Itani, 1967, p.
90).
Dorsal pelage is relatively long in M. fuscata.
Although mean length of hairs on the back in
adults is ca. 40 mm, length of the longest hairs in
this region is ca. 70-90 mm (Inagaki, 1985, p.
335; 1986, p. 117; cf. Hori et al., 1977, p. 315).
The proximal half of individual dorsal hairs is
pale gray; the distal half is marked by one to three
pairs of alternating pale and dark annulations
(Temminck, 1842, p. 9; Inagaki, 1986, p. 116;
1996, p. 92; Inagaki & Nigi, 1988b, p. 506); pale
annulations are yellow to golden, and dark an-
nulations are brown to blackish.
Early Development — Natal pelage in M. fus-
cata is unannulated, grayish brown to dark brown
(Forbes, 1894, p. 14; Pocock, 1906, p. 566; Hir-
aiwa, 1981, p. 313; Hamada et al., 1992, p. 7;
Pavelka, 1993, p. 61). Facial skin in newborns is
pale pink, paler than in adults. Replacement of the
unannulated pelage of newborns by annulated pel-
age, generally similar to that of older monkeys,
commences about age 2 months (Takada, 1966, p.
73; Maruhashi, 1982, p. 320), and this replace-
ment is complete by age 4-5 months.
Seasonal Variation— M/M5cato undergoes an
annual molt in late spring/early summer (Table 3).
In nine immature and adult captives maintained
outdoors at the Japan Monkey Centre, Inuyama,
Aichi Prefecture, molting occurred from April to
August, and the duration of the molting process
FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES
Fig. 3. Macaca fuscata — adult male, adult female, and infant — photographed 13 March 2002 at Takasakiyama,
Oita Prefecture, Kyushu. (Photo courtesy of Minoru Kinoshita, PRIKU.)
in individual monkeys varied from less than 1
month to approximately 4 months. In all five par-
turient females in this group, molting occurred af-
ter parturition.
As a result of the molting cycle, dorsal pelage
is relatively short and dark during the summer
(distal dark annulations exposed, proximal pale
gray region not yet fully exposed) and relatively
long and pale during the winter (Mori, 1979b, p.
373; Inagaki and Nigi, 1988a, p. 83; Hamada et
al., 1992, p. 19). During late winter/early spring,
the annulated tips of the hairs become progres-
sively abraded, and the pelage appears faded and
scraggly.
Geographic Variation — Thomas (1906
["1905"], p. 361) and Kuroda (1940, p. 273) ap-
parently were the first authors to indicate that dor-
sal pelage color in M. fuscata tends to be paler in
the northern part of its range than in the southern
part. Although few localized museum study skins
are available to document this cline (see Appen-
dix 1), requisite data have been provided by Ha-
mada et al. (1992, p. 1), who conducted an inten-
sive photometric study of living monkeys of
known geographic origin (Table 2). Hamada et al.
(1992, p. 9) have shown that, in females and
males from Yakushima, Kojima (= Koshima),
Kyushu (Takasakiyama), and Honshu, paleness of
dorsal pelage color in M. fuscata generally in-
creases with increasing latitude and decreasing
winter temperature; three museum skins collected
on Shikoku (BM(NH) 1906.1.4.1-1906.1.4.3) are
relatively dark and therefore suggest that pelage
color on this island also is compatible with the
cline documented by Hamada et al. (cf. Thomas,
1906 ["1905"], p. 361). As noted by Hamada et
FIELDIANA: ZOOLOGY
Table 2. Geographic variation in mean paleness index values of pelage on dorsal surface of trunk, dorsal surface
of hands, and dorsal surface of feet in Macaca fuscata; greater index values indicate paler pelage.'
Locality
of group
Latitude
(N)
Longitude
(E)
Mean
January
tempera-
ture (»C)
Mean paleness
; index values-
[or original
locality
of group
progenitors]'
Females
Males
Hands,
Back dorsum^
Feet,
dorsum**
Back
Hands,
dorsum^
Feet,
dorsum**
Honshu
Shiga
Nikko
[Wakasa]
[Takagoyama]
[Arashiyama]
Hagachi
36°44'
36°44'
35°20'
35° 13'
35°00'
34°4r
138°26'
139°30'
134°24'
139°59'
135°4r
138°44'
-4.7
-4.9
1.85
3.7
2.8
5.6
39 45(1.15)
37 41(1.11)
32 38(1.19)
38 40(1.05)
33 39(1.18)
32 31 (0.97)
Kyushu
50(1.28)
45(1.22)
43(1.34)
48(1.26)
46(1.39)
37(1.16)
37
33
29
37
29
30
42(1.14)
37(1.12)
38(1.31)
37(1.00)
34(1.17)
32(1.07)
47(1.27)
42(1.27)
45(1.22)
42(1.45)
37(1.23)
Takasakiyama
33°10'
131°27'
4.9
30 31 (1.03)
Kojima
36(1.20)
27
27(1.00)
32(1.19)
Kojima
31°27'
131°23'
6.7
31 31(1.00)
Yakushima
36(1.16)
29
32(1.10)
35(1.21)
[Yakushima*]
Nagata
30°34'
30°34'
130°26'
130°26'
11.2
11.2
31 27(0.87)
31 27(0.87)
33(1.06)
29 (0.94)
29
25
24 (0.83)
23 (0.92)
35(1.21)
26(1.04)
' Reference: Hamada et al. (1992, pp. 4, 10).
- Standard deviation, extremes, and sample size not published in cited reference.
' Brackets indicate original localities of progenitors of four sampled groups that had been translocated one or two
generations previous to the cited study, as follows: [Wakasa] and [Arashiyama] group progenitors translocated to
PRIKU, Inuyama, Aichi Prefecture; [Takagoyama] group progenitors translocated to Higashi-Tsukuba, Utopia Zoo,
Yasato-cho, Ibaraki Prefecture; and [Yakushima] group progenitors translocated to JMC, Ohirayama, Inuyama, Aichi
Prefecture (cf. Kawai, 1960, p. 183).
■* Hands/back paleness index ratio in parentheses.
^ Feet/back paleness index ratio in parentheses.
^ Hamada et al. (1992, p. 3) suspect that pelage color in this translocated group (Yakushima to Inuyama) may have
changed in response to its new environmental conditions.
Table 3. Months during which molting (M) was observed in nine Macaca fuscata captives' maintained outdoors
at the Japan Monkey Centre, Inuyama, Aichi Prefecture, Honshu; for parturient females, the month of parturition (P)
also is indicated. -
Age
(yr)
Sex
Spec. no. March
April
May
June
July
August
2
$
58
3
6
46
5
9
3
5
9
4
>6
9
43
>6
9
38
>6
9
73
>6
3
5
>6
S
42
M
M
M
M
M
M
P
M
M
P
M
M
M
M
P
M
M
P/M
M
M
M
M
M
M
Origin: Kyoto Prefecture, Honshu.
Reference: Inagaki and Nigi (1988a, pp. 82, 83).
FOODEN AND AI1V1I: SYSTEMATIC REVIEW OF JAPANESE MACAQUES
Table 4. Dorsal hair density (number of hairs per 1 -cm-diameter circle) in infant and adult Macaca fuscata:
studv conducted on open-air groups during winter 1983-1984.'
Locality
Latitude
(N)
Longitude
(E)
Mean
annual
tempera- .
ture (°C)
Hair density-
Infants
Adults
Island
Mean ± SD
N
Mean ± SD
N
Honshu
Honshu
Mivajima
Shimokita'
Hagachi-*
Miyajima-cho^
41-08'
34-~4r
34-17'
140-50'
138-44'
132-18'
9.1'
ca. 16"
ca. 16^^
1625.0 = 134.4
1116.3 r 202.8
3
0
1028.9 = 309.4
793.5 ± 96.6
745.0 i 226.4
12
4
4
■ Reference: Inagaki and Hamada (1985. p. 87).
- Data for females and males combined: no significant difference between sexes.
' Studied in November 1983.
■•Studied in December 1983.
^ Studied in January 1984: progenitors of this group were translocated from Shodoshima (ca. 34''30'N. 134-18'E)
to Miyajima-cho in 1962.
al. (1992. p. 18). the pigmentation cline in M. fus-
cata accords well with Gloger's rule (cf. Mayr.
1963. p. 324).
Neonatal pelage color apparently follows a sim-
ilar latitudinal cline (Hamada et al.. 1992. p. 7:
cf. Kuroda. 1984. p. 14). In four captive neonates
(age <4 months) that originated on Yakushima.
the mean dorsal pelage paleness index \ alue was
27.9. and in five captive neonates that originated
on Honshu, this value was 30.8.
Pelage on the dorsal surface of the hands and
feet also generally a\erages paler with increasing
latitude and decreasing winter temperature on
Yakushima. Kojima. Kyushu (Takasakiyama), and
Honshu (Table 2); at all localities studied, the dor-
sal surface of the feet averages paler than the dor-
sal surface of the hands in both sexes. The dorsal
surface of the hands is particularly dark (blackish)
in Yakushima M. fuscata. On Kojima. Kyushu.
Shikoku (BMfNHJ 1906.1.4.1-3). and Honshu,
the dorsal surface of the hands averages paler
than, or approximately as pale as. the dorsal sur-
face of the trunk (hands/back paleness index ratio.
0.97-1.31); on Yakushima. the dorsal surface of
the hands averages distinctly darker than the dor-
sal surface of the trunk (hands/back paleness in-
dex ratio. 0.83-0.92). Available infonnation per-
mits no inference concerning whether variation of
the hands/back paleness index ratio in the Yaku-
shima population of M. fuscata is completely dis-
junct from that in non- Yakushima populations of
this species.
Latitudinal variation of dorsal hair density in
M. fitscata has been studied at three localities b\
Inagaki and Hamada (1985. p. 85). In accord with
thermoregulatory expectations, adult hair density
at these localities increases with increasing lati-
tude and decreasing mean annual temperature (Ta-
ble 4: cf. Hori et al.. 1977. p. 316). Hair density
in infants (age <1 year), which has been studied
at two of these localities, also apparently increases
with increasing latitude and decreasing mean an-
nual temperature; this suggests that the thermo-
regulatory adaptation is genetic, not merely phys-
iological.
Albinism — Albino M. fitscata individuals have
been captured in Tochigi and Yamagata Prefec-
tures, northern Honshu (Temminck. 1842. p. 10;
Kuroda. 1940. p. 271). Al Takasakiyama. Oita
Prefecture. Kyushu, during the period 1950-1963.
Itani et al. (1964. p. 30; cf. Itani & Mizuhara.
1957. p. 106) observed abnormal, sharply defined
patches of white fur. usually on the hands and
rarely on the feet, in 27 monkeys (10 females. 17
males) of a total population of 771 individuals; on
the hands, the smallest of these white patches
were restricted to the fingertips, whereas the larg-
est extended as far as the distal forearms (cf. Tro-
isi et al.. 1981. p. 6. who report patches of white
fur on the fingers in one of 54 monkeys in a cap-
tive group translocated from Takasakiyama to It-
aly). Similar white patches also have been report-
ed on the limbs of "more than six" monkeys at
Hagachi. Shizuoka Prefecture. Honshu, and on the
limbs of "a few" monkeys on Kojima. off the
east coast of Kyushu (Nishida. 1966. p. 199; cf.
Yamagiwa. 1979. p. 492).
External Measurements and Proportions
LiNE.AR Measurements of Body Size — The
standard mammalogical measurement of body
size is head and bodv length, which is the distance
FIELDIANA: ZOOLOGY
Table 5.
Fig. 4).'-2
Prefectural variation in head and body length and related external measurements in Macaca fuscata (cf.
Mean
coordinates
Adult females
Adult males
Head and
Relative
Body
Head and
Relative
Body
of sample
body length
tail length
weight
body length
tail length
weight
Prefecture
localities
(mm)
(T/HB X 100)
(kg)
(mm)
(T/HB X 100)
(kg)
Honshu
Niigata
38°18'N
532.0
15.4
10.1
561
16.0
12.2
139°36'E
U)
(/)
{1)
{1)
{1)
{1)
Nagano
35°58'N
528.2 ± 25.82
15.2 ± 3.08
8.9 ± 1.51
580.1 ± 33.08
14.9 ± 4.14
12.6 ± 2.17
137°55'E
477-572
4.4-20.8
5.5-13.8
499-630
3.5-21.7
8.8-18.4
{42)
{42)
{42)
{31)
{30)
{31)
Shimane
34°52'N
488.5 ± 20.80
17.8 ± 3.25
7.9 ± 1.33^
511.9 ± 27.93
16.2 ± 4.17
8.6 ± 1.17
132°38'E
464-517
13.0-22.8
6.0-9.2
472-545
8.8-19.9
7.6-10.8
{6)
(6)
(5)
(7)
(7)
(7)
Mie^
34°20'N
136°30'E
—
—
—
650
(i)
13.8
(i)
—
Shikoku
Tokushima
33°50'N
515.8 ± 58.68
16.3 ± 2.21
8.2
—
—
—
134°18'E
472-601
{4)
13.3-18.6
{4)
7.8-8.6
(2)
Kochi
33°24'N
—
—
—
597
12.4
11.2
133°03'E
U)
(i)
{1)
Totals
—
—
522.8 ± 30.50
15.6 ± 3.09
8.8 ± 1.49
570.1 ± 42.30
15.1 ± 3.99
11.8 ± 2.49
464-601
4.4-22.8
5.5-13.8
472-650
3.5-21.7
7.6-18.4
{53)
{53)
{50)
{41)
{40)
{40)
' Within each cell, figures indicate mean ± SD (where n > 2), extremes, and sample size (italicized, in parentheses).
^ This table excludes measurements of ""Head and body length" reported by Napier and Napier (1967, p. 406),
who cite "Iwamoto et al. (1966) personal communication" as source; the published measurements actually are
measurements of sitting height, not head and body length (M. Iwamoto, letter, 3 Feb. 1998). Also excluded are two
measurements reported as head and body length by Aizawa and Hagiwara (2001, p. 5); these are measurements of
anterior trunk length (M. Hagiwara, pers. comm., 3 Oct. 2002).
' Excludes one pregnant adult female, body weight = 9.4 kg.
4 Data from Kuroda (1940, p. 271).
from the tip of the nose to the base of the tail
(Imaizumi, 1970, p. 89). This measurement is
available for 94 wild-collected adult specimens of
M. fuscata — 53 females collected in three prefec-
tures on Honshu and one prefecture on Shikoku
and 41 males collected in four prefectures on
Honshu and one prefecture on Shikoku (Table 5);
nearly 80% of these specimens were collected in
Nagano Prefecture, Honshu. Mean (±SD) head
and body length is 522.8 ± 30.50 mm (extremes,
464-601 mm) in the available female sample and
570.1 ± 42.30 mm (extremes, 472-650 mm) in
the available male sample. Although the unbal-
anced geographic distribution of the available
samples hinders inference concerning geographic
variation of head and body length in M. fuscata,
this measurement apparently averages less in both
sexes in Shimane Prefecture, southwestern Hon-
shu, than in the more northeasterly prefectures on
Honshu for which data are available (Table 5; Fig.
4).
Sitting height, the distance from the apex of the
crown to the plane of the ischial callosities
(Schultz, 1929, pp. 218, 233), is an anthropolog-
ical measurement of body size that is similar but
not identical to head and body length; this mea-
surement is available for 168 adult females (mean
= 537.4 mm) and 126 adult males (mean = 590.7
mm) that originated on five Japanese islands (Ta-
ble 6). Inter-island comparison indicates that sit-
ting height in M. fuscata is smallest on Yakushi-
ma, as previously noted by Iwamoto (1971, p.
154); somewhat greater on Shodoshima; and
greatest on Kyushu and Honshu (Fig. 5; Table 6);
evidence provided by the small Shikoku sample
is equivocal. On Honshu, no general pattern of
latitudinal or longitudinal variation in sitting
height is evident (Appendix 5); however, this
FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES
650
600
550
500
450
• •
•
•
•
•
Adult males
o
•
•
•
•
• •
•
•
•
•
•
•
0
•
600
0 - - ^
<■
•
550
••:.
•
Adult females
o
•
•
* ••
•
•
•
500
«
• •
• ^
o
(
, •
•
•
•
• Honshu
o Shikoku
450
33
34
35
36
Latitude ("N)
37
38
39
Fig. 4. Latitudinal variation in head and body length in Honshu and Shikoku samples of Macaca fuscata adults
(cf. Table 5).
measurement apparently averages less on Boso
Peninsula (Appendix 5: 35-36°N. 139-14rE)
than elsewhere on Honshu (Boso Peninsula — fe-
males, 525.7 mm, n = 6; males, 579.5 mm, n =
1 1; elsewhere on Honshu — females, 552.1 mm, n
= 95; males, 604.6 mm, n = 58), and it also may
be relatively small in males in southwestern Hon-
shu (south of 35°N, west of 134°E; Appendix 5).
The linear measurement of body size in M. fus-
cata for which the largest samples are available is
anterior trunk length, an anthropological measure-
ment of the distance from the suprasternal notch to
the cranial margin of the pubic symphysis (Schultz,
1929, pp. 232, 234); this measurement is available
for 783 adult females (mean = 365.2 mm) and 350
adult males (mean = 392.5 mm) that originated on
FIELDIANA: ZOOLOGY
Table 6. Insular variation in sitting height (mm)
in Macaco
fuscata adi
Lilts (cf. Fig. 5).'
Adult females
Adult males
Mean latitude
Island
of localities (N)
Mean ± SD
N
Mean ± SD
N
Honshu-'
35°37'
555.5
± 29.21^
75
602.5
± 34.395
61
Honshu-
34°30'
536.3
± NA*
26
586.1
± NA"
8
Shodoshima^
34°30'
518.8
± 14.47
20
574.9
± 22.58
21
Shikoku^
33°34'
527.5**
2
609
1
Kyushu^
33°10'
537.1
± 19.08
11
621.2
± 19.54
8
Yakushima^
30°34'
510.1
± NA*
34
568.1
± NA*
27
Totals
—
537.4
± NA"
168
590.7
± NA"
126
' Reference: Iwamoto (1971, table opp. p. 156), except as otherwise indicated in footnote 3.
^ For details concerning these samples, see Appendix 5.
-'' Data in this row were derived from specimens examined, not from Iwamoto (1971).
'' Extremes: 487-625 mm.
5 Extremes: 504-682 mm.
^ NA = not available.
' Captive and/or provisioned populations.
** Extremes: 512-543.
seven Japanese islands (Table 7). In accord with
Bergmann's rule (cf. Mayr, 1963, p. 320), anterior
trunk length generally increases with latitude from
Yakushima and Kojima to Honshu (Fig. 6), as pre-
viously noted by Hamada et al. (1996a, p. 106).
As indicated above for sitting height, no general
pattern of latitudinal or longitudinal variation in an-
terior trunk length is evident in Honshu samples
(Appendix 6). Also as indicated for sitting height,
anterior trunk length apparently averages less on
Boso Peninsula (Appendix 6, 35-36°N, 139-
14 TE) than elsewhere on Honshu (Boso Peninsu-
la— females, 351.5 mm, n = 23; males, 394.7 mm,
n = 16; elsewhere on Honshu — females, 374.7
mm, n = 338; males, 403.1 mm, n = 194), and
anterior trunk length tends to be relatively small in
a small sample of adult males collected in south-
western Honshu (south of 35°N, west of 133°E;
Appendix 6).
Tail Length — Mean tail length in M. fuscata
is 82.6 mm in 665 adult females and 91.1 mm in
312 adult males (Table 8). The tail in both sexes
tends to be relatively long on Shodoshima and
Yakushima (Table 8, Fig. 8), as previously noted
by Hamada et al. (1996a, p. 105), and it tends to
be relatively short on Kojima and probably on
Shikoku. The apparent difference in tail length be-
tween the Yakushima and Kojima samples is per-
haps unanticipated because the distance between
these two small islands is less than 140 km.
Among females on Honshu, tail length may tend
to decrease with latitude (Fig. 7; Appendix 7).
The ratio of tail length to head and body length
(T/HB) provides a useful index value for com-
paring individuals or species of different body siz-
es (cf. Fooden & Albrecht, 1999, p. 433). In avail-
able samples collected on Honshu and Shikoku,
the mean value of T/HB is similar in adults of
both sexes — 15.6% in 53 females and 15.1% in
40 males (Table 5); this ratio averages greater in
young immatures (T/HB = ca. 25%) (Fig. 8),
which indicates that the postnatal growth rate of
head and body length exceeds that of tail length.
The ratio of tail length to anterior trunk length
(T/ATL), which is available for samples that are
much larger and geographically more broadly dis-
persed than those for T/HB, provides a supple-
mentary basis for appraisal of insular variation of
tail length relative to body size (Table 8; Appen-
dix 7). Judging from these samples, T/ATL tends
to be relatively large in both sexes on Yakushima
and Shodoshima and tends to be relatively small
in both sexes on Shikoku. Although T/ATL in the
total sample of females (22.7%) is close to T/ATL
in the total sample of males (23.3%), this ratio in
males exceeds that in females in seven of the
eight available insular samples (Table 8).
The tail is unusually short (<50 mm, T/HB
<10%) in three adult females and three adult
males collected in Nagano Prefecture, Honshu,
and in one adult male collected in Shimane Pre-
fecture, Honshu (Table 9; Fig. 8; cf. Table 8);
three of these short-tailed specimens (one female,
two males) make up the entire sample that was
collected at Shimashima, Nagano Prefecture.
Monkeys with similarly short tails (reportedly
<30 mm) have been observed but not measured
at two additional M. fuscata localities (Itani et al..
FOODEN AND AlMl: SYSTEMATIC REVIEW OF JAPANESE MACAQUES
11
700
• Honshu
D Kyushu
•
•
m
__ ..__ _
o Shodoshima
A Yakushima
D
*
" • 1
• •• • • • 1
o
•
• V
1
■
1' ^
( 1
1 «
•
•
•• •
•
550
X
• ; • •
•
?
— 500
o
•
•
Ajdult males
c
5
in
•
600
• •
• •
s •
550
•
•
•
•
•
i
-
I
o
1
o
( 1
(
•
•
•
500
J
i- •'
• 1
!
\dii\t females
450
! 1
30
32
33
34
35
36
37
38
39
Latitude (°N)
Fig. 5. Latitudinal variation in sitting height in Macaca fuscata adults (cf. Table 6). Data points with standard-
deviation error bars are mean values derived from Iwamoto (1971, table opp. p. 156); data points without error bars
are individual values derived from specimens examined (except for one male data point with coordinates x = 34.47
and y = 595, which is an individual value from Iwamoto, 1971).
1964, p. 35): at Takasakiyama, Kyushu, seven ab-
normally short-tailed adults or subadults (three fe-
males, four males) were seen in a total population
of 771, and on Kojima, off the eastern coast of
Kyushu, one abnormally short-tailed adult male
was seen in a total population of ca. 50 (cf. Ka-
wai, 1963, p. 113).
Hind Foot Length — Mean hind foot length in
Macaca fuscata is 153.6 min in 657 adult females
and 170.4 mm in 273 adult males (Table 10). Hind
foot length generally tends to increase with lati-
tude from Yakushima to Honshu (Fig. 9). How-
ever, in female and male samples available from
Shodoshima, hind foot length is notably less than
expected for the latitude of this island. In Honshu
samples, hind foot length is not closely correlated
with latitude (Appendix 8).
The ratio of hind foot length to anterior trunk
length (HF/ATL), which has been reported for
several M. fuscata samples by Hamada et al.
12
FIELDIANA: ZOOLOGY
Table 7. Insular variation in anterior trunk length (mm) in Macaca fiiscata adults (cf. Fig. 6).'
Adult females
Adult males
Mean latitude
Island
of localities (N)
Mean ± SD
N
Mean ± SD
N
Honshu-"*
35°43'
376.6 ± 23.71"
88
398.2 ± 28.975
68
Honshu-
35°37'
373.3 ± NA*
273
404.6 ± NA"
142^
Shodoshima**
34°30'
355.0 ± 19.48
19
392.8 ± 21.93
20
Awajishima
34°14'
353.3 ± 8.03
9
387.4 ± 21.12
19
Kyushu**
33°10'
363.7 ± 18.03
278
392.4 ± 23.70
22
Shikoku
33°04'
347.3 ± NA^
179
398
110
Kojima"
31°27'
346.2 ±20.13
55
366.0 ± 26.89
43
Yakushima^
30°34'
338.7 ± 15.84
44
367.8 ± 22.83
35
Totals
—
365.2 ± NA^
783
392.5 ± NA"
350
' Reference: Hamada et al. (1996a, pp. 101, 102), except as otherwise indicated in footnotes 3, 7, 9, and 10.
^ For details concerning these samples, see Appendix 6.
-^ Data in this row were derived from specimens examined, not from Hamada et al. (1996a).
4 Extremes: 329-470 m.
■''Extremes: 315-450 mm.
* NA = not available.
^ Includes two measurements (400 mm, 404 mm) that were originally reported as head and body length by Aizawa
and Hagiwara (2001, pp. 5-6); see Table 5, footnote 2.
^ Captive and/or provisioned populations.
' Includes two specimens examined (anterior trunk length 337 and 360 mm).
'° Specimen examined.
" Semiprovisioned population (ca. 12 kg of wheat distributed weekly to ca. 70 monkeys).
(1996a, pp. 104, 105), provides a useful index val-
ue that facilitates comparison of hind foot pro-
portions in individuals of varying body size. As
noted above for hind foot length, HF/ATL is no-
tably low in the Shodoshima female and male
samples; this ratio is notably high in the Kojima
female and male samples. Although mean HF/
ATL in the total sample of females (42.2%) is
close to that in males (43.1%), this ratio in males
exceeds that in females in all seven available pairs
of insular samples (Table 10) (cf. above. Tail
Length).
Body Weight — Body weight in M. fuscata av-
erages 8.4 kg in 832 adult females and 1 1.3 kg in
368 adult males (Table 11; cf. Hazama, 1964, p.
94; Hiraiwa, 1981, p. 314). As previously noted
by Hamada et al. (1996a, p. 100), body weight in
this species generally tends to increase with lati-
tude (Fig. 10), in accord with Bergmann's rule;
however, a small sample of two low-weight adult
males (5.6 kg and 6.5 kg) collected on Kinkazan,
an islet off northeastern Honshu, is a conspicuous
exception to this generalization. Body weight also
averages exceptionally small in 36 adult females
(7.0 kg) and 25 adult males (9.5 kg) studied on
Boso Peninsula, southeastern Honshu (Appendix
9, 35-36°N, 139-14rE).
Seasonal changes in body weight have been re-
ported in M. fuscata. At Shiga and Hakusan,
which are relatively cool habitats, body weight
tends to decrease by up to ca. 2 kg during the
winter (Hamada et al., 1996b, p. 320; cf. Hiraiwa,
1981, p. 314). Conversely, on Kojima, a warm
habitat, body weight tends to decrease by approx-
imately 2 kg during the summer (Mori, 1979b, p.
387; Horii et al., 1982, p. 427). In a group of
laboratory captives of unknown origin that was
housed in an air-conditioned room and fed a stan-
dardized diet, mean body weight decreased by ca.
1 .5 kg during the winter months (Matsubayashi &
Enomoto, 1983, p. 523; cf. Hamada et al., 2003,
p. 85), approximately as in free-ranging monkeys
living in cool habitats (see above).
Translocation of M. fuscata to cooler or warmer
climates apparently modifies body weight in a
manner predicted by Bergmann's rule (Paterson,
1996, p. 589). Twenty-seven years after translo-
cation of a group from Mihara, Hiroshima Prefec-
ture, to a cooler climate in Oregon (U.S.A.), fe-
male and male growth curves for body weight in
the descendant group had become significantly
higher than those in the original group. Converse-
ly, 20 years after translocation from Arashiyama,
Kyoto Prefecture, to a warmer climate in Texas
(U.S.A.), growth curves for body weight in the
descendant group had become lower than those in
the original group.
Summary — Available data concerning sitting
FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES
13
475
450
425
400
375
350
325
!
.0.
I
■ Awajishima
• Honshu
A Kojima
p Kyushu
o Shikoku
* Shodoshima
A Yakushima
—
1
j
•■
Adult males r
1 .J.
tI
r4
*•
•
i
1
[
r 1 P 'r
1 .11 J
1 1
i
' 1
j
-
—
J
i
L
i^
• •
• 1 j
! j
■
»
i i
1
!
^
• '
■ill
300
30
31
32
33
34
35
36
Latitude (°N)
37
38
39
40
41
42
Fig. 6. Latitudinal variation in anterior trunk length in Maccica fuscata adults (cf. Table 7). Data points with
standard-deviation error bars are mean values derived from Hamada et al. (1996a, pp. 101-102); data points without
error bars are individual values derived from specimens examined.
height, anterior trunk length, and body weight in-
dicate that body size in M. fuscata generally tends
to increase with latitude from Yakushima to Hon-
shu, in accord with Bergmann's rule. On Honshu,
sitting height and anterior trunk length generally
do not increase with latitude, but body weight
does. Data concerning sitting height, anterior
trunk length, and body weight all indicate that
body size is unusually small on Boso Peninsula,
southeastern Honshu. Body size probably also is
unusually small on Kinkazan island, off north-
eastern Honshu, but this inference is based exclu-
sively on body weight data available for two adult
males.
14
FIELDIANA: ZOOLOGY
Table 8. Insular variation in tail length (mm) and tail length/anterior trunk length ratio (T/ATL) in Macaco
fuscata adults (cf. Figs. 7-8).'
Mean
latitude of "
localities _
(N)
Adult females
Adult males
Tail length
Mean ± SD
N
T/ATL
Tail length
Island
Mean ± SD
N
T/ATL
Honshu-''
35°5r
81.7 ± 15.00
54
21.7%
88.3 ± 19.79
49
22.2%
Honshu^
35°37'
79.3 ± NA^
213
21.2%
91.1 ± NA-»
125=
22.5%
Shodoshima^
34°30'
87.2 ± 9.14
19
24.6%
102.1 ± 9.24
21
26.0%
Awajishima
34° 14'
77.3 ± 3.81
9
21.9%
88.0 ± 8.65
19
22.7%
Shikoku
33°ir
73.5 ± NA4
17^
20.8%
74
1«
18.6%
Kyushu""
33°10'
86.5 ± 11.16
260
23.8%
94.0 ± 14.76
21
24.0%
Kojima'o
3r27'
73.3 ± 8.69
49
21.2%
80.4 ± 12.77
41
22.0%
Yakushima"
30°34'
90.0 ± 8.25
44
26.6%
101.4 ± 10.61
35
27.6%
Totals
—
82.6 ± NA^
665
22.7%
91.1 ± NA^
312
23.3%
' Reference: Hamada et al. (1996a, pp. 101-102, 104-105), except as otherwise indicated in footnotes 2, 5, 7,
and 8.
- Specimens examined.
'' For details concerning these samples, see Appendix 7.
■* NA = not available.
^ Includes one specimen reported by Aizawa and Hagiwara (2001, p. 5; cf. Table 5, footnote 2); cf. Kuroda (1940,
p. 271).
'' Captive and/or provisioned populations.
'' Includes two specimens examined (cf. Table 7).
•* Specimen examined (cf. Table 7).
** For tail length measurements in a Macaca fuscata group that was translocated from Kyushu (Takasakiyama) to
Italy, see Masali and Scarsini (1981, p. 12).
'" Semiprovisioned population (ca. 12 kg of wheat distributed weekly to ca. 70 monkeys).
Tail length and T/ATL, which are exceptionally
high on Yakushima and Shodoshima and excep-
tionally low on Kojima, apparently are not closely
correlated with latitude. Conversely, hind foot
length, like body size, tends to increase with lat-
itude in M. fuscata from Yakushima to Honshu,
but it does not tend to increase with latitude on
Honshu. Hind foot length and HF/ATL are nota-
bly low on Shodoshima.
Dermatoglyphics
Dermatoglyphic patterns on the hands of M.
fuscata were studied by Iwamoto and Suryobroto
(1994, p. 78; cf. Iwamoto, 1964, p. 54; 1975a, p.
73; De Stefano et al., 1981b, p. 34; Iwamoto &
Suryobroto, 1990, p. 433) in five samples that
originated on Honshu, Kyushu (two samples), Ko-
jima, and Yakushima. Although no absolute dif-
ferences in palmar patterns or palmar ridge direc-
tions were found among these samples, several
frequency differences were discovered in these
traits (Table 12); frequencies in the Shimokita
Peninsula (Honshu) and Yakushima samples, re-
spectively, at the northern and southern limits of
distribution of M. fuscata, were the most deviant
among the five samples studied. On the feet of M.
fuscata, frequency differences in plantar derma-
toglyphic patterns have been discovered in Taka-
sakiyama (Kyushu) and Yakushima samples (Iwa-
moto, 1967b, p. 156; cf. De Stefano et al., 1981a,
p. 22).
Intraspecific variation in palmar dermatoglyph-
ic pattern frequencies within M. fuscata apparent-
ly may exceed interspecific variation in pattern
frequencies among M. fuscata, M. fascicularis,
and M. nemestrina (Cauble & Mavalwala, 1973,
p. 151; Morris & Kerr, 1974, p. 229).
Congenital Malformation of Limbs
In 1954 and 1955, two or three years after reg-
ular provisioning of several free-ranging M. fus-
cata groups had been initiated, primatologists in
Japan began to notice the occurrence of monkeys
with congenitally malformed appendages (Itani &
Mizuhara, 1957, p. 106; Itani et al., 1964, p. 30;
Furuya, 1966, p. 488; Shidei et al., 1981, p. 1).
The severity of observed malformations varied
from syndactyly, to deformed interphalangeal
FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES
15
130
120
110
100
90
80
70
60
50
40
I 30
£" 20
o
= 110
n
I-
100
90
80
70
60
50
40
30
20
■r *
i
■ Awajishima
• Honshu
A Kojima
D Kyushu
o Shikoi<u
o Shodoshima
r ',
T
»•
•
1 ■ r
1
• >
i 1
i :
j
II 1 -l-T
'"J-. •
1.
1
1
i ir
•
;
A Yakushima
1
!
o
■
J
'•
i
' 1
1 !
!
i
• i
•
1
i
'
•
i
j
i
1 ; j
•
!
Adult males
i
«
•
■
. lu
d'-
1
' , .1
> T T
II
1)
li
•
<•
•
1
,
<
»
( 1
1
1 1
j
[^ J
i
■ 1
" !•
!■•■•; *
•
»
■'■
1
1
1
■■
1 !
i i
!
• !
* • !
; 1
Adult females
•
30
34
36
Latitude (°N)
37
38
39
40
42
Fig. 7. Latitudinal variation in tail length in Macaca fuscata adults (cf. Table 8). Data points with standard-deviation
error bars are mean values derived from Hamada et al. (1996a, pp. 101-102); data points without error bars are
individual values derived from specimens examined or from literature records cited in Table 8.
joints, to missing digits (usually middle digits) on
hands and/or feet, to absence of hands or feet, to
virtually complete absence of pelvic limbs; only
one instance of Polydactyly has been reported
(Homma, 1980, p. 26).
The geographic distribution of M. fuscata
groups with malformed members extends broadly
from Shiga, north-central Honshu, to Yakushima,
at the southern limit of the specific range (Itani et
al., 1964, p. 30; Furuya, 1966, p. 489; Nishida,
1966, p. 199; Iwamoto, 1967b, p. 248; Tanaka &
Nigi, 1967, pp. 93, 96; Iwamoto & Hirai, 1970,
p. 395; Iwamoto et al., 1975, p. 78; Yoshihiro et
al., 1979, p. 459; Homma, 1980, p. 21; Shidei et
al., 1981, p. 17; Nakamichi et al., 1983, p. 53;
Minezawa et al., 1990, p. 573). Of 65 groups sur-
veyed for congenital limb malformations, 22
(33.8%) included malformed individuals (Table
16
FIELDIANA: ZOOLOGY
Table 9. External measurements in seven Macaco
fuscata adults with unusually short tails (cf. Figs. 7-8).
Head
and
body
Tail
PRIKU
length length
Locality field no. Sex
(mm)
(mm)
T/HB
Nagano Prefecture
!, Honshu
Aoki 87 ?
529
46
8.7%
Nishiharuchika 120 9
505
42
8.3%
Shimashima 39 9
502
22
4.4%
Kamimachi 41 6
600
21
3.5%
Shimashima 38 6
537
45
8.4%
Shimashima 40 S
557
38
6.8%
Shimane Prefecture, Honshu
Ushiroyama (H-9906) 6
545
48
8.8%
pie autosomal dominant or as a simple autosomal
recessive (Iwamoto, 1967b, p. 261; Iwamoto &
Hirai, 1970, p. 396; Yoshihiro et al., 1979, p. 468;
Shidei et al., 1981, p. 28; Minezawa et al., 1990,
p. 573). Inbreeding, pesticide exposure, and con-
tamination of provisioned food have all been sug-
gested as possible etiological factors (Nishida,
1966, p. 199; Yoshihiro et al., 1979, p. 469; Shidei
et al., 1981, p. 35; Minezawa, 1990, p. 575). Al-
though elevated organochlorine pesticide residues
have been detected in cadavers of malformed
monkeys and in cadavers of mothers of mal-
formed monkeys, this evidence is not yet conclu-
sive as to the cause of malformations (Ito et al.,
1988, pp. 109, 111, 112).
13). The occurrence of malformed monkeys in
provisioned groups appears to be more common
than in nonprovisioned groups.
The incidence of congenital limb malforma-
tions in newborn infants apparently varies locally
and varies over time at a single locality. At nine
localities where affected groups have been studied
for 2-26 years, the mean neonatal incidence of
malformations varies from ca. 2.0% to 17.3% (Ta-
ble 14); at one of these localities (Awajishima),
the annual incidence varied from 71.4% in 1971
to 0% in 1972 to ca. 40% in 1977 (Yoshihiro et
al., 1979, p. 461). The incidence of malformations
in female infants apparently is similar to that in
male infants (Table 15); results of one study in-
dicate that primiparous mothers are less likely to
produce malformed infants than are multiparous
mothers (Nakamichi et al., 1997, p. 229). During
the 1980s, the incidence of malformations is said
to have declined rapidly (Minezawa et al., 1990,
p. 575).
The cause or causes of the congenital limb mal-
formations are unclear (Iwamoto, 1967b, p. 261;
Morris, 1971, p. 338; Yoshihiro et al., 1979, p.
469; Shidei et al., 1981, p. 39; Ito et al., 1988, p.
1 12; Minezawa et al., 1990, p. 576; Nakamichi et
al., 1997, p. 231). Some mothers with malforma-
tions have produced infants with malformations,
and some affected individuals are known to be
siblings (Itani et al., 1964, p. 34; Iwamoto, 1967b,
p. 248; Nakamichi et al., 1997, p. 232; cf. Hase-
gawa & Iwasaki, 1984, p. 399), which may sug-
gest a genetic influence. However, malformations
are not associated with detectible karyological ab-
normalities, and pedigree analysis tends to rule
out inheritance of malformations either as a sim-
Cranial Characters (Figs. 11-14)
E^ect of Provisioning
Some free-ranging groups of M. fuscata — such
as those inhabiting Kojima, Shodoshima, Taka-
goyama, and Takasakiyama — are artificially pro-
visioned to varying degrees (see Gazetteer, Ap-
pendix 4). To investigate the effect of artificial
provisioning on skull size, greatest length of skull
in a sample of provisioned adults collected at Ta-
kagoyama, Chiba Prefecture, has been compared
with that in a sample of nonprovisioned adults
collected elsewhere in the same prefecture (Table
16). Mean greatest length of skull in the provi-
sioned females (1 18.1 mm) and males (134.4 mm)
significantly exceeds that in the nonprovisioned
females (115.8 mm) and males (129.5 mm). For
this reason, where provisioned samples are in-
cluded in subsequent analyses, they are individu-
ally identified. Data from skulls of monkeys
raised in captivity are excluded from subsequent
analyses.
Sex and Age Variation
In the available sample of nonprovisioned wild-
collected adult specimens of M. fuscata, greatest
length of the skull in males (133.2 ± 5.72 mm, n
= 100) averages about 12% greater than in fe-
males (119.1 ± 4.94 mm, n = 141) (Table 17);
size differences between M. fuscata males and fe-
males reportedly are already evident in the skulls
of early juveniles (Mouri, 1994, p. 46). Although
rostral-postrostral ratio (XlOO) in adult males
FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES
17
in
CM
o in o
CM T- T-
'\ •*• of ■ •
• tv
. , o '.
• . • ■
• . •. •
1 1 '. •• . ■ ■
o Females, adult
• Males, adult
o Females, immature or ac
• Males, immature or age
o
in
CD
o
o
CO
o
in
in
o
ti
o
r^
in
^
,^*^
.—
E
:=
E
>
C
JC
~~-
■*^
^
o
O)
:;
u;
-^
c
1
>x
r-
•a
•~
o
—
^
tl
"O
JJ
o
o
c
re
_>1
•D
^
re
-o
o
X
ij
o
,^-
in
>
CO
o
o
o
in
o
o
CO
(lUiu) i{;6u3| ijei
FIELDIANA: ZOOLOGY
Table 10. Insular variation in hind foot length (mm) and hind foot length/anterior trunk length ratio (HF/ATL)
in Macaca fuscata adults (cf. Fig. 9).'
Mean
Adult females
Adult males
latitude of
localities _
(N)
Hind foot
Mean ± SD
length
N
HF/ATL
Hind foot
length
Island
Mean ± SD
N
HF/ATL
Honshu-
35°37'
159.0 ± NA^
238
42.5%
177.3 ± NA-^
135^
44.0%
Shodoshima^
34°30'
139.0 ± 4.45
19
39.2%
159.6 ± 5.37
21
40.6%
Awajishima
34° 14'
147.6 ± 5.01
8
41.8%
168.6 ± 4.98
18
43.5%
Kyushu'^
33°10'
155.1 ± 5.80
276
42.6%
170.3 ± 8.25
21
43.4%
Shikoku
33°06'
150.8 ± NA^
176
43.4%
—
0
—
Kojima'
31°27'
152.6 ± 6.01
55
44.1%
164.1 ± 8.21
43
44.8%
Yakushima^
30°34'
137.0 ± 7.03
44
40.5%
155.7 ± 6.48
35
42.3%
Totals
—
153.6 ± NA-^
657
42.5%
170.4 ± NA^
273
43.1%
' Reference: Hamada et al. (1996a, pp. 101-103), except as otherwise indicated in footnotes 4 and 6.
^ For details concerning these samples, see Appendix 8.
^ NA = not available.
■* Includes one specimen reported by Kuroda (1940, p. 271) and one specimen reported by Aizawa and Hagiwara
(2001, p. 5; cf. Table 5, footnote 2); the specimen reported by Kuroda lacks ATL information and therefore is not
included in the HF/ATL calculation.
'^ Captive and/or provisioned populations.
^ Includes two BM(NH) specimens examined; these specimens lack ATL information and therefore are not included
in the HF/ATL calculation.
'' Semiprovisioned population (ca. 12 kg of wheat distributed weekly to ca. 70 monkeys).
(57.8 ± 3.65. n = 100) averages about 13% great-
er than in adult females (51.2 ± 3.03, n = 137),
relative zygomatic breadth (XlOO) in males (7L7
± 2.22, n = 98) is approximately the same as that
in females (70.9 ± 2.16, n = 139). From infancy
to adulthood, rostral-postrostral ratio in males in-
creases 105%, whereas relative zygomatic breadth
increases only 10%.
Dental emergence norms in M. fuscata are pre-
sented in Table 18. For further details concerning
sex and age variation in skulls of this species, see
Schweyer (1909, p. 12), Ikeda and Watanabe
(1966, p. 272), Koike and Shimamura (1988, p.
77), and Mouri (1994, p. 46; 1996, p. 283).
Geographic Variation
Judging from available data, mean greatest
length of skull in M. fuscata increases successive-
ly from south to north on Yakushima, Kojima, and
Kyushu, the three southernmost islands represent-
ed by samples (Table 19; Fig. 15); this is in accord
with Bergmann's rule (cf. Mayr, 1963, p. 320).
Farther northward, however, mean skull length
apparently does not increase from south to north
on Shikoku, Shodoshima, Honshu, and Kinkazan;
in fact, mean skull length apparently decreases on
Kinkazan, an islet off the northeastern coast of
Honshu. Intra-island variation of skull length in
the relatively large sample of Honshu specimens
is analyzed in Appendix 10; as previously noted
by Kuroda (1989, p. 7), the most conspicuous fea-
ture of this variation is the aberrantly small skull
size in females and males collected on Boso Pen-
insula, southeastern Honshu (Appendix 10, cell
35-36°N, 140-141°E; cf. Fig. 15).
A somewhat related series of measurements of
"head length" (greatest midsagittal length from
glabella to occiput) in living M. fuscata from sev-
en islands has been published by Hamada et al.
(1996a, pp. 102-104) (Table 20); the head length
measurement employed in this study, which is
standard in anthropology, excludes the rostrum of
the skull and therefore pertains only to the cra-
nium. The study of Hamada et al. further differs
from the present one in including some specimens
from captive and/or translocated groups. Despite
these methodological differences, the pattern of
insular variation indicated by measurement of
head length has many similarities to that indicated
by measurement of greatest length of skull (cf.
Tables 19, 20).
Ontogenetic development of greatest length of
skull in Yakushima M. fuscata is compared with
that in Honshu M. fuscata in Figure 1 6. A mean
size difference between specimens from these two
islands is already apparent in skulls with newly
FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES
19
lyo
190
185
180
175
170
Adult males
■ Awajishima
• Honshu
A Kojima
D Kyushu
o Shikoku
o Shodoshima
A Yakushima
- -
J
M.
••
(
1
( )
n
fr
■
(
( )
- -
- r"'
II
1
■'■
( 1
■tRe^
1
1Rn
' T'' ^
k
■■
■'■
i
I
i
155
—
■1-
1) 145
iJ 175
o
•2 170
■o
c
I 165
160
lies
T
Ad
ult femj
r
(
1
(
■
•
1 1
•
I
1
i<
1
i
( 1
■
••
^'^^
1
I
1
J-
(
■-
J
i
145
140
135
130
125
[
C
6
0
II
T
( 1
I
i
■■
■■
■•
■■
I
<'
"" i
]
' i
30
31
32
33
34
35
36
Latitude (°N)
37
38
39
40
41
42
Fig. 9. Latitudinal variation in hind foot length in Macaca fuscata adults (cf. Table 10). Data points with standard-
deviation error bars are mean values derived from Hamada et al. (1996a, pp. 101), data points without error bars are
individual values derived from specimens examined or from literature records cited in Table 10.
erupted third molars (age ca. 6 years); available
evidence is inadequate to determine the age of
onset of this size difference.
Ikeda and Watanabe (1966, p. 274; cf. Kuroda,
1984, p. 15) have shown that orbital index (orbital
height/orbital breadth) averages greater in M. fus-
cata collected on Yakushima than in M. fuscata
collected on other Japanese islands (cf. Table 21).
The average insular difference in this index is pri-
marily the result of lesser orbital breadth in Yak-
ushima M. fuscata — not of greater orbital height
(Fig. 17).
Computed tomographic imaging of a sample of
46 M. fuscata skulls that originated at six locali-
ties— extending from Yakushima to Shimokita
Peninsula — has revealed that nasal cavity area in
this species tends to increase with increasing lat-
itude, whereas maxillary sinus volume tends to
decrease with increasing latitude (Rae et al., 2003,
p. 155). Enlargement of the nasal cavity at higher
latitudes has been interpreted by Rae et al. as an
adaptation that functions to augment the warming
of inspired air in cooler climates; reduction of the
maxillary sinuses at higher latitudes is interpreted
20
FIELDIANA: ZOOLOGY
Table 11. Insular variation in body weight (kg) in Macaca fuscata adults (cf. Fig. 10).'
Mean
Adult females
Adult males
latitude of
localities (N)
Island
Mean ± SD
N
Mean ± SD
N
Kinkazan-
38°17'
_
0
6.0^
2
Honshu"*
35°54'
9.6 ± NA^
276
12.9 ± NA5
143^
Honshu^"
35°38'
8.7 ± 1.56^
125
11.4 ± 2.44»
81
Shodoshima^
34°30'
8.9 ± 1.15
18
10.7 ± 2.73
20
Awajishima
34°14'
7.1 ± 1.26
10
10.8 ± 1.58
19
Kyushu'^
33°10'
7.8 ± 1.10
281'«
10.8 ± 2.15"
22
Shikoku
33°05'
8.2 ± NA5
1712
11.2
1'^
Kojima'^
31°27'
6.3 ± 0.70
55
8.0 ± 1.83
43
Yakushima*
30°34'
8.0 ± 1.60
44
10.2 ± 2.06
35
Yakushima-
30°20'
5.3 ± 0.95'5
616
9.5'^
29
Totals
—
8.4 ± NA'
832
11.3 ± NA^
368
' Reference: Hamada et al. (1996a, pp. 101, 102), except as otherwise noted in footnotes 2, 6, 10, 12, and 13.
^ Data in this row were derived from specimens examined, not from Hamada et al. (1996a).
^ Individual values: 5.6 kg, 6.5 kg.
•* For details concerning these samples, see Appendix 9.
^ NA = not available.
6 Includes two specimens recorded by Aizawa and Hagiwara (2001, p. 5).
^ Extremes: 4.8-13.8 kg; includes one known pregnant female, weight = 9.4 kg.
^Extremes: 7.6-18.4 kg.
' Captive and/or provisioned individuals.
'" Includes one specimen examined, 6.3 kg.
"Cf. Nigi et al. (1980, p. 236).
'- Includes two specimens examined (see Table 5).
''' Specimen examined (see Table 5).
'* Semiprovisioned population (ca. 12 kg of wheat distributed weekly to ca. 70 monkeys).
'^ Extremes: 4.0-6.5 kg.
'* Includes four captives.
'^ Individual values: 9.2 kg, 9.8 kg.
as a passive result of crowding by the adjacent
enlarged nasal cavity.
Based on frequency of occurrence of selected
nonmetric cranial characters, Kuroda (2002b, p.
125) has reported that skulls of the Boso Penin-
sula, Shodoshima, and Yakushima populations of
M. fuscata tend to differ from skulls of other stud-
ied populations of this species; relevant frequen-
cies are not specified in this report. In another
publication, Kuroda (2002a, p. 115) has compared
metric cranial characters of M. fuscata subfossils,
dated Early Jomon to Late Jomon (ca. 10,000-
300 B.C.), with those of modern populations from
various Japanese localities; discriminant functions
indicate that measurements of most of the subfos-
sils are generally similar to those of modern pop-
ulations.
Dental morphology and measurements in M.
fuscata skulls originating on Yakushima are com-
pared in detail with those in skulls originating on
other unspecified Japanese islands by Saheki
(1966, p. 412). In another dental study, Yoshika-
wa and Deguchi (1992, p. 123) found that per-
manent maxillary canines were double-rooted in
9 of 22 female M. fuscata specimens examined
(40.9%) and in 0 of 22 male specimens (all single-
rooted); collecting localities of the specimens
were not reported. Dental enamel thickness in M.
fuscata has been studied by Shimizu (2002, p.
613).
Interspecific Comparisons
M. fuscata has the largest skull and the most
projecting rostrum of any of the four species in
the fascicularis group (Table 22; cf. Ehara, 1974,
p. 92). Development of a well-defined median
sagittal crest is relatively uncommon in adult
males of M. fuscata (23 of 158 specimens ex-
amined; cf. Ikeda & Watanabe, 1966, p. 275), as
in M. mulatta and M. cyclopis (Fooden, 2000, p.
44; Fooden & Wu, 2001, p. 9).
FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES
21
18
16
12
i 4
O 14
OQ
12
•
Adult
males
•
i*
(, • T •
T «
T' T-"
„{■
^ 1l
1
T
T
i-J
lb
•
o
II
<>
w ■>
• •
■ Awajishima
• Honshu
♦ Kinkazan
A Kojima
D Kyushu
o Shikoku
o Shodoshima
A Yakushima
—
A
■■
-1-
•
•
<
••
4
• .
B
•
♦
j
♦
j I
Adult f
emales
•
1
•
1
•
•
»
-•I
•
tv *
•
<
1
( 1
T
i" ^t!'
ft ^
1
I
^Z
.
V
iS- o ^ Jl
II
D i
r
!
• i
i
A
•
•
1
1
i
30
35
36
Latitude (°N)
Fig. 10. Latitudinal variation in body weight in Macaca fuscata adults (cf. Table 11). Data points with standard-
deviation error bars are mean values derived from Hamada et al. (1996a, pp. 101-102); data points without error
bars are individual values derived from specimens examined or from literature records cited in Table 1 1 .
Molecular Biology and Genetics
DNA: Intraspecific Studies
Mitochondrial DNA — The most important
studies of individual and local variation in
mtDNA in M. fuscata have been published by
Hayasaka et al. (1986, p. 348; 1991, p. 401) and
Kawamoto (1997, p. 32; 1998, p. 53; 1999, p.
302; 2002, p. 61; cf. Hayaishi & Kawamoto,
2002, p. 164). Hayasaka et al. sequenced the ma-
jor noncoding region of mtDNA in a sample of
100 monkeys; Kawamoto analyzed mtDNA re-
striction site patterns in a composite sample of
134 monkeys, sequenced 412 base pairs in the D-
loop in a sample of 119 monkeys, and (with co-
author Hayaishi) sequenced 203 base pairs in the
D-loop in another sample of 38 monkeys. Local-
ities sampled Kawamoto include those sampled
by Hayasaka et al.
22
FIELDIANA: ZOOLOGY
Table 12. Frequency (%) of dermatoglyphic patterns on both hands in five local samples of Macaca fuscata.^
Localities
Takasaki-
Shimokita,
Kawaradake,
yama,
Dermatoglyphic
Pattern
Honshu
Kyushu
Kyushu
Kojima
Yakushima
pattern
type
(N = 70)
(N = 106)
(N = 236)
(N = 234)
(N = 276)
Thenar
O
11.4
63.7
59.6
67.2
22.5
L
88.6**
36.3
40.0
32.8
77.5**
W
0
0
0.4
0
0
First interdigital
O
5.7
0
1.3
0
0
L
92.9
92.1
68.4
100.0
80.7
W
1.4
7.9
30.3**
0
19.32
Second interdigital
L
21.4**
2.0
0.9
5.6
1.1
W
78.6
98.0
99.1
94.4
98.9
Third interdigital
O
1.4
3.0
2.6
0
0.7
L
17.4
37.6^
19.7
49.1**
13.5
W
81.2
59.4
77.7
50.9
85.8
Fourth interdigital
L
1.4
0
1.3
4.7
8.0
W
98.6
100.0
98.7
95.3
92.0
Distal hypothenar
o
71.4
61.0
69.9
75.6
100.0**
L
28.6
37.0
23.3
21.4
0
W
0
2.0
6.8
3.0
0
Proximal hypothenar
L
1.4
37.8
35.7
47.6
97.1**
W
98.6**
62.2
64.3
52.4
2.9
Ending of ridge
Prox
0
85.1
82.7
91.4
5.1
direction II
Furc
0
5.0
6.9
0.9
1.8
Ulna
100.0**
9.9
10.3
7.7
92.7**
X
0
0
0
0
0.4
Ending of ridge
Prox
0
10.0
8.6
3.9
1.1
direction III
Furc
0
3.0
3.0
1.7
0
Ulna
100.0
81.0
82.8
92.7
95.6
Other
0
6.0
5.6
1.7
3.3
' Reference: Iwamoto and Suryobroto (1994, pp. 78, 82). Double asterisks (**) indicate values that are significantly
different from others in the same row (P < 0.01), except as otherwise noted.
- Not significantly different from Takasakiyama value (P > 0.05).
^ Not significantly different from Kojima value (P > 0.05).
Table 13. Provisioning status and occurrence of
congenital limb malformations in 65 Macaco fuscata
groups.'
Provisioning
status of group
Groups with
malformed members
Number %
. Number
of groups
Free ranging
Nonprovisioned
Provisioned
5
14
Captive
23.8
40.0
21
35
Provisioned
3
33.3
9
Totals
22
33.8
65
' Reference: Yoshihiro et al. (1979, p. 460); cf. Shidei
et al. (1981. p. 16).
Hayasaka et al.'s (1986, p. 348; 1991, p. 401)
sample of 100 monkeys originated at 12 localities
that span entire geographic range of the species
from northernmost Honshu to Yakushima (Fig.
18). Sequencing of the major noncoding region
revealed that eight mtDNA haplotypes (designat-
ed types I-VIII) are present in these 100 samples
(Table 23). Based on sequence similarities, three
of the eight mtDNA haplotypes may be allocated
to group II/IA^II, three other haplotypes may be
allocated to group III/IVA^III, and the remaining
two haplotypes (V and VI) are of uncertain rela-
tionships.
In group \y\fW\\, mtDNA haplotype II includes
one copy of a repeatable 158-bp sequence, hap-
lotype I includes two copies of the same se-
quence, and haplotype VII includes three copies
(Hayasaka et al., 1991, p. 406). Similarly, in
group III/IV/VIII, mtDNA haplotypes III, IV, and
VIII, respectively, include one, two and three cop-
FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES
23
Table 14. Neonatal incidence of congenital limb malformations in nine Macaca fuscata groups.'
Incidence of
congenital
limb malformations
Observation
Locality
Island
period
Number
%
N
Provisioned groups
Shiga
Honshu
1963-1977
19
12.4
153
Otoumi
Honshu
1972-1974
2
5.4
37
Ohirayama
Honshu-
-1956-1966
3
-2.0
-150
Gagyusan
Honshu
1955-1977
66
12.2
542
Hagachizaki
Honshu
1972-1975
8
12.1
66
Miyajima-cho
Miyajima^
1961-1978
17
9.0
188
Awajishima
Awajishima
1969-1995
132
17.3
762
Takasakiyama
Kyushu
1965-1978
Captive group
126
3.7
3395
Takaosan
Honshu"*
1970-1977
2
8.0
25
Totals
—
—
375
-7.1
-5318
' References: Iwamoto (1967b, p. 249; Ohirayama group only); Yoshihiro et al. (1979, p. 460); Nakamichi et al.
(1997, p. 227; Awajishima group only); cf. Shidei et al. (1981, p. 16).
- Group studied after translocation from Yakushima to Ohirayama.
^ Group studied after translocation from Shodoshima to Miyajima.
"* Group progenitors captured at Hasumi (Honshu) and Shodoshima.
ies of a repeatable 167/168-bp sequence. Haplo-
types V and VI do not include repeated sequenc-
es. Haplotypes I and VII probably were derived
from haplotype II by segmental duplication and
triplication, respectively (Hayasaka et al., 1991, p.
411), and haplotypes IV and VIII probably were
similarly derived from haplotype III. In a se-
quenced 896-bp HindWl fragment, the number of
pairwise nucleotide differences reported within
group II/I/VII is nine, and the number of differ-
ences reported between group II/IA^II and group
III/IVA^III is 14-17 (Hayasaka et al., 1996, p.
1048).
Of the 1 2 localities studied, nine are represent-
ed by multiple specimens (2-32 specimens), and
three are represented by one specimen each (Table
23). Mitochondrial DNA haplotypes are mono-
Table 15. Sexual variation in incidence of congen-
ital limb malformations in Macaca fuscata neonates.'
Incidence of congenital
limb malformations
Sex
Number
%
N
Females
Males
Totals
71
95
166
A.V
6.02
5.3
1520
1585
3105
' Reference: Yoshihiro et al. (1979, p. 462); cf. Shidei
et al. (1981, p. 30); Nakamichi et al. (1997, p. 227).
2P = 0.10.
morphic at each of six localities represented by
multiple specimens, and they are polymorphic at
the remaining three localities represented by mul-
tiple specimens; because sample size averages
smaller at monomorphic localities (6.5 specimens
per locality; extremes, 2-15) than at polymorphic
localities (19.3 specimens per locality; extremes,
8-32), it is possible that larger sample sizes would
reveal a higher incidence of local mtDNA hap-
lotype polymorphism. At one of the polymorphic
localities (Nikko), seven specimens are haplotype
II, and one is haplotype VI (dimorphism); these
two haplotypes are not members of the same hap-
lotype group. At the other two polymorphic lo-
calities, most specimens included two distinguish-
able mtDNA haplotypes (heteroplasmy)-haplo-
types 1 and VII (both members of group II/I/VII)
at Takahama and haplotypes IV and VIII (both
members of group III/IV/VIII) on Yakushima.
Judging from available data, the most common
and widespread mtDNA haplotype in M. fuscata
is haplotype II, a member of group II/I/VII. Hap-
lotype II occurs at seven localities over a span of
ca. 900 km, from Shimokita Peninsula in northern
Honshu to Wakasa (Tottori Prefecture) in west-
central Honshu and to Shodoshima, an islet in the
Inland Sea off the southern coast of west-central
Honshu. At Takahama, which is within the area
of distribution of haplotype II, 12 of 18 specimens
are heteroplasmic for haplotypes I and VII; as in-
dicated above, haplotypes I and VII probably
24
FIELDIANA: ZOOLOGY
were derived from haplotype II by segmental du-
plication and triplication. Also within the area of
distribution of haplotype II is Nikko, where seven
of eight specimens are haplotype II and one is
haplotype VI, and Izu, where the single available
specimen is haplotype V; the relationship of hap-
lotype VI and haplotype V to other haplotypes is
uncertain.
Limited data available from three southern lo-
calities suggest that the III/IVA^III group of
mtDNA haplotypes replaces the II/I/VII group
west and south of Wakasa and Shodoshima. At
Gagyusan (Okayama Prefecture), which is on
Honshu and is only 60-90 km west of Wakasa
and Shodoshima, all five specimens examined are
mtDNA haplotype III. And on Yakushima, which
is ca. 570 km southwest of Gagyusan, all 32 spec-
imens examined are either haplotype IV or hap-
lotype VIII or both (heteroplasmic); haplotypes
IV and VIII probably were derived from haplo-
type III by segmental duplication and triplication.
However, at Takasakiyama, which is on Kyushu
and is between Gagyusan and Yakushima, all
three specimens examined are haplotype V, a
mtDNA haplotype of uncertain relationships, oth-
erwise known in M. fuscata only in the single
specimen from Izu, which is on Honshu ca. 700
km east-northeast of Takasakiyama.
In summary, the most important systematic im-
plications of Hayasaka et al.'s (1991, p. 406) se-
quencing study of the major noncoding region of
mtDNA in 100 M. fuscata samples representing
12 Japanese localities are as follows:
1. Eight mtDNA sequence haplotypes (I-VIII)
are known in M. fuscata.
2. Mitochondrial DNA haplotypes I and VII
probably were derived from mtDNA haplotype II
by segmental duplication and triplication, respec-
tively (group II/I/VII). Similarly, mtDNA haplo-
types IV and VIII probably were derived from
mtDNA haplotype III by segmental duplication
and triplication (group III/IV/VIII). The relation-
ship of haplotype V and haplotype VI to other
haplotypes is uncertain.
3. Mitochondrial DNA haplotype is monomor-
phic at six of nine Japanese localities represented
by multiple samples, it is dimorphic at one local-
ity, and it is heteroplasmic at two localities.
4. Mitochondrial DNA haplotype II occurs at
7 of 12 sampled localities, and mtDNA haplotype
V occurs at two localities. Each of six other
mtDNA haplotypes occurs at only one locality.
5. The II/I/VII group of haplotypes is wide-
spread in northern and central Honshu, and this
group also occurs on nearby Shodoshima.
6. The III/IV/VIII group apparently replaces
the II/I/VII group in southwestern Honshu and on
Yakushima.
7. Mitochondrial DNA haplotype V is disjunct-
ly distributed in south-central Honshu and Kyu-
shu.
8. Mitochondrial DNA haplotype VI is known
only in one of eight specimens obtained at Nikko,
north-central Honshu.
Kawamoto's (1997, p. 32; 1998, p. 53; 1999, p.
302) restriction-site studies yielded results that are
largely compatible with results of his geographi-
cally more comprehensive sequencing study
(2002, p. 65). For this reason, only the sequencing
study will be discussed here.
Kawamoto's (2002, p. 57) sequencing study of
119 monkeys included samples collected at 106
localities spanning the entire geographic range of
M. fuscata, from Shimokita Peninsula in the north
to Yakushima in the south. Of 49 haplotypes de-
tected in this study, 31 were restricted to one lo-
cality, six occurred at two localities, four at three
localities, three at four localities, one at five lo-
calities, two at seven localities, one at 12 locali-
ties, and one at 18 localities (Table 24, Fig. 19).
Localities that share the same haplotype tend to
be fairly tightly clustered geographically; how-
ever, this does not apply to the 18 localities that
share haplotype no. 1, which span ca. 400 km in
northern Honshu, and also does not apply to the
seven localities that share haplotype no. 20, which
span ca. 400 km in central Honshu; unlike hap-
lotype no. 20, haplotype no. 1 appears to be the
only haplotype present in most of its broad area
of distributions. Haplotypes are polymorphic at 10
of 1 1 localities represented by multiple samples
(monomorphic at one locality).
Based on sequence divergence values, with M.
cyclopis taken as an out-group, the 49 sequence
haplotypes that Kawamoto (2002, p. 65) detected
in M. fuscata are clearly separable into two
groups — group I, 17 haplotypes (nos. 7-11, 35-
43 [including 38 + 1, 41 - 1, 41 + 1]), mean
pairwise difference = 8.81 ± 4.19 (SD), and
group II, 32 haplotypes (nos. 1-6, 12-23, 26-34
[including 16 + 1, 17 + 1, 17 + 2, 20 + 1, 26
+ 1]), mean pairwise difference = 5.98 ± 2.88.
In groups I and II combined, with a total of 49
haplotypes, mean pairwise difference = 9.10 ±
4.22.
Although the known geographic distribution of
group I haplotypes is mainly in southern Japan
FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES
25
Fig. 11. Skull of adult female Macaca fuscata fuscata-
Honshu. Scale in cm. (Photographs by Minoru Kinoshita.)
-PRIKU 6492, Kuradama. Kimitsu. Chiba Prefecture,
(Fig. 19), extending from Yakushima to Awajish-
ima and southwestern Honshu (Gagyusan, Oka-
yama Prefecture), group I haplotypes also occur
disjunctly in a relatively small area in east-central
Honshu (eight specimens in Yamanashi, Tokyo,
and Saitama Prefectures; one specimen in Niigata
Prefecture). Group II haplotypes are widely and
apparently continuously distributed from Sho-
doshima and west-central Honshu (Wakasa, Tot-
tori Prefecture) to northernmost Honshu (includ-
ing Kinkazan islet). Although group II localities
virtually surround the disjunct group I localities
in east-central Honshu, group I haplotypes and
group II haplotypes are not known to occur at the
same locality.
Because group I haplotypes are more differen-
tiated than group II haplotypes, Kawamoto (2002,
p. 68) has inferred that the group I population
originated earlier than the group II population.
Kawamoto (2002, p. 69) has further suggested
26
FIELDIANA: ZOOLOGY
Fig. 12. Skull of adult female Macaca fuscata yakui — PRIKU 6638, Hanyama, Kamiyaku-cho, Kagoshima Pre-
fecture, Yakushima. Scale in cm. (Photographs by Minoru Kinoshita.)
that the divergence of group I and group II pop-
ulations and the broad dispersal of haplotype no.
1 (group II) in northern Honshu are consequences
of changes in climate and vegetation that were
induced by Pleistocene glacial advance and re-
treat.
In a supplementary publication, Hayaishi and
Kawamoto (2002, p. 164) reported on D-loop se-
quence haplotypes (203 bp) in 38 M. fuscata sam-
ples that were collected at 21 localities widely dis-
persed on Yakushima. Analysis of these samples
revealed four haplotypes (Y1-Y4) that differed at
only three of 203 sites. Thirty-three samples col-
lected at 18 localities were identified as Yl (cf.
Domingo-Roura et al., 2004, p. 37); two samples
collected at two close-lying localities were iden-
tified as Y2; two samples collected at one locality
were identified as Y3; and one sample — collected
at a locality that also yielded a sample identified
as Yl — was identified as Y4.
FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES
27
Fig. 13. Skull of adult male Macaca fuscata fiiscata-
Scale in cm. (Photographs by Minora Kinoshita.)
-PRIKU 5414. Naka, Natashou, Fukui Prefecture, Honshu.
Following is a brief summary of the results of
Kawamoto's (2002, p. 61) sequencing study of
119 A/, fiiscata samples representing 106 locali-
ties from Shimokita Peninsula to Yakushinia:
1. Of 49 haplotypes detected. 31 were restrict-
ed to a single locality.
2. Haplotypes shared by multiple localities
tend to cluster in local areas. However, each of
two exceptional haplotypes (nos. 1 and 20) is dis-
tributed over a span of ca. 400 km; haplotype no.
1 apparently is the only haplotyjje present in most
of its broad area of distribution.
28
FIELDIANA: ZOOLOGY
Fig. 14. Skull of adult male Macaca Juscata yakiii — PRIKU 1571, Hanyama, Kamiyaku-cho, Kagoshima Prefecture,
Yakushima. Scale in cm. (Photographs by Minoru Kinoshita.)
3. Haplotypes are polymorphic at 10 of 1 1 lo-
calities represented by multiple specimens.
4. Sequence divergence separates the 49 de-
tected haplotypes into two groups — group 1 (17
haplotypes) and group II (32 haplotypes).
5. Group I haplotypes are distributed from
Yakushima to southwestern Honshu and disjunct-
ly in east-central Honshu; group II haplotypes are
continuously distributed from west-central Hon-
shu to northernmost Honshu. Group I haplotypes
and group II haplotypes are not known to occur
together at the same locality.
FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES
29
Table 16. Greatest length of skull (mm) in sample of provisioned Macaca fuscata population collected at Tak-
agoyama,' Chiba Prefecture, Honshu, compared with that in sample of nonprovisioned populations collected elsewhere
in Chiba Prefecture.
Sample statistics
Provisioned
Nonprovisioned
Mean ± SD
Extremes
N
Mean ± SD
Extremes
N
118.1 ± 2.85
113.7-125.5
20
134.4 ± 3.26
128.6-138.1
Adult females
115.8 ± 3.35
109.2-120.9
25
Adult males
129.5 ± 4.22
121.3-139.0
21
2.44
2.95
<0.022
<0.012
' Cf. Koike and Shimamura (1988, p. 75).
- Two-tailed test.
-* Excludes one Takagoyama specimen, collected ca. 1961, for which provisioning history is unknown (JMC 2588,
greatest length of skull = 134.1).
6. Greater genetic differentiation of group I
haplotypes suggests that the group I population
may be older than the group II population.
7. Divergence of group I and group II popu-
lations and broad dispersal of haplotype no. 1
(group II) may be related to Pleistocene glacial
advance and retreat.
The most salient point of agreement in the re-
sults published by Hayasaka et al. (1986, p. 348;
1991, p. 401) and Kawamoto (1997, p. 32; 1998,
p. 53; 1999, p. 302; 2002, p. 61) is that, based on
sequence data, mtDNA haplotypes in M. fuscata
are broadly subdivisible into two geographic
groups — a northeastern group, distributed in
northern and central Honshu, and a mainly south-
western group, distributed primarily in western
Table 17. Cranial measurements' and proportions in age/sex classes of nonprovisioned wild-collected Macaca
fuscata.-
Greatest length
Relative zygomatic
Postrostral
Rostral-postrostral
of skull
breadth
length
ratio
Age/sex class"*
(mm)
(ZB/GL X 100)
(mm)
(R/PR X 100)
Infants
90.4 ± 5.14
65.1 ± 1.98
75.9 ± 3.82
28.2 ± 2.70
77.5-104.4
60.1-69.1
66.5-86.2
22.3-33.2
{51)
{51)
(52)
(57)
Juveniles
106.8 ± 9.57
68.7 ± 2.19
82.9 ± 4.62
40.6 ± 6.23
86.9-132.8
63.3-76.4
72.1-98.1
28.7-58.3
{283)
{280)
{284)
{282)
Subadult females
113.5 ± 3.99
69.9 ± 1.79
83.9 ± 2.96
A1.6 ± 1.83
107.2-121.4
67.1-73.9
78.1-88.2
44.0-52.4
{27)
(26)
{27)
{27)
Subadult males
128.9 ± 6.03
69.7 ± 2.05
91.6 ± 3.85
54.3 ± 2.71
113.8-143.1
66.2-73.8
82.9-100.0
49.0-61.1
{40)
{39)
{39)
{39)
Adult females^
119.1 ± 4.94
70.9 ± 2.16
86.5 ±3.18
51.2 ± 3.03
109.2-133.7
64.3-77.9
79.1-94.8
43.5-57.5
U41)
{139)
{138)
{137)
Adult male.s^
133.2 ± 5.72
71.7 ± 2.22
92.0 ± 3.05
57.8 ± 3.65
120.6-148.3
65.5-77.3
83.0-98.8
48.3-66.2
{100)
{98)
{105)
{100)
' For definition of measurements, see Fooden (1969, p. 40).
- Mean ± SD, extremes, and sample size (italicized figures in parentheses).
' Dental specifications: infants, deciduous teeth only; juveniles, some permanent teeth erupted; subadults, M3 in
females or C in males incompletely erupted; adults, all permanent teeth completely erupted.
"Cf. Fig. 15.
30
FIELDIANA: ZOOLOGY
Table 18. Dental emergence-'' chronology in Macaca fuscata. (References: Iwamoto et al., 1984, p. 277; 1987,
p. 24; cf. Nass, 1977, p. 309; Passarello, 1980, p. 54).
Tooth
Sex
Dental emergence age (yr)
Dispersion (ca. 10th percentile-
Mean ca. 90th percentile)
h
m,, m'
c,, c'
nij
m2
M,
M'
I2
P
M2
M2
C,
P3. P'
C
P,, P'
P4, P4
c,
c
M3, M-^
F + M
F + M
F + M
F + M
F + M
F + M
F + M
F + M
F + M
F + M
F + M
F + M
F + M
F + M
F
F
M
F + M
M
M
F + M
Deciduous teeth*
0.019
0.000-0.038
0.058
0.019-0.096
0.077
0.038-0.12
0.21
(0.15-0.27)^
0.21
(0.15-0.29)^
0.46
(0.38-0.56)^
0.50
(0.42-0.60)7
Permanent teeth
8
1.50
1.00-2.00
1.75
1.25-2.25
2.50
2.00-3.00
2.75
2.25-3.25
3.25
2.75-3.75
3.50
3.00-4.00
3.75
3.25-4.25
3.75
3.25-4.50
4.00
3.50-4.50
4.00
3.50-4.75
4.25
3.75-4.75
4.25
3.75-4.75
4.50
4.00-5.25
4.75
4.25-5.50
5.75
5.00-6.75
39 + 37
39 + 37
39 + 37
39 + 37
39 + 37
39 + 37
39 + 37
101 + 106
101 + 106
101 + 106
101 + 106
101 + 106
101 + 106
101 + 106
101
101
101
106
101 + 106
106
106
101 + 106
' "* Tooth position indicators.
^ Defined as initial penetration of gingiva by crown of tooth.
* Samples include 31 captives (16 females, 15 males) and 45 free-ranging monkeys (23 females, 22 males).
'' Dispersion values reported for these teeth are extreme emergence ages.
^ Samples include 39 captives (17 females, 22 males) and 168 free-ranging monkeys (84 females, 84 males). Samples
of free-ranging populations at Shiga (31 females, 24 males) and Kojima (46 females, 35 males) were excluded from
calculation of the tabulated statistics by Iwamoto et al. (1987, pp. 19, 24); dental emergence ages in these two
populations apparently are ca. 25-45% greater than in other populations of M. fuscata.
Table 19. Insular variation in greatest length of skull (mm) in wild-collected Macaca fuscata (cf. Fig. 15).
Mean
latitude
of
localities .
(N)
Adult females
Adult males
Island
Mean ± SD
Extremes
N
Mean ± SD
Extremes
N
Kinkazan
38°17'
118.1 ± 1.74
114.8-119.8
7
133.9 ± 2.18
131.4-137.1
7
Honshu'
35°35'
121.0 ± 4.64
109.2-133.7
152
134.7 ± 5.25
121.3-147.7
74
Shodoshima^
34°31'
—
—
0
139.8 ± 3.23
134.4-146.3
9
Shikoku
34°00'
122.6
119.7-125.4
2
—
—
0
Kyushu^
32°53'
121.3 ± 1.20
119.6-122.4
4
142.5 ± 4.92
134.9-148.3
5
Kojima"*
31°27'
119.1 ± 4.71
109.1-129.2
19
135.2 ± 5.88
122.6-148.3
13
Yakushima
30°20'
113.4 ± 2.44
109.6-118.5
20
128.9 ± 5.23
116.2-137.4
29
Totals
—
120.0 ± 4.90
109.1-133.7
204
134.1 ± 5.99
116.2-148.3
137
' See Appendix 10.
' Provisioned population.
^ Includes three females and four males provisioned at Takasakiyama.
'' Semiprovisioned population (ca. 12 kg of wheat grain distributed weekly to ca. 70 monkeys).
FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES
31
150
145
140
135
130
125
E
E.
3 120
® 135
130
125
120
115
110
105
—
▲ D
•
!
Adult males
B
I-**- *
i
A
AA
4
A
A
A
A
A
▲
A
* • • • •
; 1
1 1
i 1
A
'<
♦
t
1 •
A
j
1
! • 1% i
i i
i i
A
•
•
i
i
!
: i i 1 1 ^
i 1 ! i :
i 1
•
•
Adult females
I " i
1
^ i
•
i
B
1 :
9
• Honshu
A
A
A
▲
A
'
1
1 ♦
■ ♦ Kinkazan
1 A Kojima
A
▲
: •
D Kyushu
' o Shikoku
▲
■,9 :
! I i
A Yakushima
30
31
32
33
34
38
39
40
35 36 37
Latitude (°N)
Fig. 15. Latitudinal variation in greatest length of skull in Macaca fuscata adults (cf. Table 19).
41
42
Honshu. Shikoku. Kyushu, and Yakushima (Figs.
18, 19). The zone of transition between these two
groups apparently is in west-central Honshu, be-
tween Wakasa (35°20'N. 134''24'E) and Gagyusan
(34°47'N. 133°37'E). Haplotypes on the islet Sho-
doshima (ca. 34°30'N, 134°15'E) apparently be-
long to the northeastern group, and haplotypes on
the islet Awajishima (ca. 34°15'N. 134°50'E) ap-
parently belong to the southwestern group.
In a recent study, Marmi et al. (2004, p. 677)
sequenced a 392-bp fragment of the mtDNA con-
trol region in samples obtained from 50 M. fiis-
cata specimens that originated at six localities in
central and southern Japan (Hakusan. n = 8; Tak-
ahama. 9; Awajishima, 8: Takasakiyama, 6; Ko-
jima. 8; Yakushima, 11); these sequences were
compared with corresponding sequences — ob-
tained from the GenBank/EMBL database — of 89
eastern M. mulatta specimens (82 from six local-
ities in China; seven from one locality in Viet-
32
FIELDIANA: ZOOLOGY
Table 20. Insular variation in head length (greatest midsagittal length from glabella to occiput, mm) in living
Macaca fuscata (Hamada et al., 1996a, pp. 102-104).
Adult females
Adult males
Island
Mean ± SD'
N
Mean ± SD'
N
Honshu-
96.6 ± 4.01
238
103.8 ±4.11
130
Shodoshima
89.6 ± 2.22
19
94.6 ±3.17
21
Awajishima
96.9 ± 4.60
9
103.4 ± 3.58
19
Shikoku
92.3 ± 3.37
15
—
0
Kyushu
94.3 ± 3.34
277
99.4 ± 4.09
21
Kojima
89.2 ± 2.71
55
92.3 ± 5.04
43
Yakushima^
89.2 ± 3.12
44
96.3 ± 4.37
35
Means
94.2 ± 3.35
657
99.9 ±4.17
269
' Extreme values not published.
2 Includes captive and/or translocated groups.
nam). Mitochondrial DNA haplotypes are mono-
morphic at each of four of the M. fuscata locaH-
ties and polymorphic at the remaining two (Tak-
ahama; Kojima). The results of neighbor-joining
and maximum-likelihood analysis in this study al-
locate five of the six M. fuscata localities to two
groups (Hakusan/Takahama and Awajishima/Tak-
asakiyama/Kojima), which coincides with a major
result of the studies of Hayasaka et al. (1986, p.
348; 1991, p. 401) and Kawamoto (1997, p. 32;
1998, p. 53; 1999, p. 302; 2002, p. 61) (see
above); evidence in Marmi et al.'s study concern-
ing the group relationship of the sixth locality
(Yakushima) is equivocal. Compared with M. mu-
latta haplotypes, the number of nucleotide differ-
ences averages less in Yakushima haplotypes
(28.3 ± 2.39; extremes, 24.8-32.0) than in hap-
lotypes at o\hev M. fuscata localities (41.4 ± 4.12;
34.7-51.6).
In other research projects, mtDNA variation has
been employed in studies of hybridization, sub-
fossils, and male dispersal. Kawamoto et al.
(1999, p. 57; 2001, p. 17) used restriction-site
analysis and sequencing of the mtDNA D-loop to
compare M. fuscata, M. cyclopis, and putative hy-
brids in Wakayama Prefecture. Mouri et al. (2000,
p. 87) and Agatsuma and Iwagami (2002, p. 75)
compared the D-loop sequence in several M. fus-
cata subfossils with that in modem M. fuscata;
among their findings is the discovery that, ca. 400
years ago, M. fuscata had been transported by hu-
mans from Yakushima to Okinawa, where the
species does not naturally occur. Yoshimi and Ta-
kasaki (2003, p. 71) sequenced the D-loop in 16
monkeys in Okayama Prefecture and concluded
that two local males had dispersed >100 km from
their natal troop.
MiNISATELLITE DNA AND MiCROSATELLITE
DNA — Analysis of minisatellite DNA and micro-
satellite DNA in blood samples has been used
successfully to determine paternity in captive
groups of M. fuscata (Weiss et al., 1988, p. 75;
Inoue et al., 1990, p. 564; 1991, p. 204; Soltis et
al., 1997a, p. 739). Analysis of microsatellite
DNA is feasible in smaller and/or more degraded
samples, including samples of semen, urine, and
feces, and therefore is suitable for noninvasive de-
termination of parentage in natural groups (Inoue
& Takenaka, 1993, p. 41; Domingo-Roura et al.,
1997, p. 358; 2004, p. 37; Hayakawa & Takenaka,
1999, p. 301). Local variation in microsatellite al-
lele frequencies has been reported for two M. fus-
cata localities (Wakasa, n = 28; Arashiyama, n =
34) by Inoue & Takenaka (1993, p. 41).
DNA: Interspecific Studies
Mitochondrial DNA — Within the fascicularis
group, mtDNA in M. fuscata apparently is much
more similar to that in M. mulatta and M. cyclopis
than to that in M. fasciculahs (Table 25; cf. Mouri
et al., 2000, p. 92; Domingo-Roura et al., 2004,
p. 37). Within M. mulatta, mtDNA varies geo-
graphically (Melnick et al., 1993, p. 289; Morales
& Melnick, 1998, p. 16; Tosi et al., 2002, p. 168);
mtDNA in M. fuscata is more similar to that in
eastern M. mulatta (China, Burma) than to that in
western M. mulatta (India, Pakistan).
Nuclear DNA: Noncoding Intron — Deinard
and Smith (2001, p. 47) sequenced a fragment of
intron 4 (—410 bp) and the entire length of intron
5 (—545 bp) of the natural resistance-associated
macrophage protein 1 (NRAMPl) gene in 59
FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES
33
150
140
130
120 -
110
100
90
« 80
o 140
130
Males
8 •
t
120
110
100
90
80
Females
• • • 4
i!
• Honshu
A Yakushima
10 12
Estimated age (yr)
14
16
20
Fig. 16. Greatest length of skull vs. estimated age in Honshu and Yakushima specimens of Macaca fuscata. Age
estimates to age ca. 6 years are based on dental emergence chronology (cf. Table 18): age estimates beyond age 6
years are based on degree of tooth wear (relative to tooth wear in known-age Kojima specimens).
34
FIELDIANA: ZOOLOGY
Table 21. Insular variation in orbital measurements and orbital height/breadth index in adult wild-collected
Macaca fuscata (cf. Fig. 17).
Orbital breadth (mm)
Orbital height (mm)
Orbital index (H/Br)
Island
Mean ± SD
Extremes
Mean ± SD
Extremes
Mean ± SD
Extremes
N
Adult females
Honshu
24.4 ± 1.02
21.9-27.2
24.1 ± 1.70
20.5-28.0
0.99 ± 0.056
0.89-1.14
88
Shodoshima
—
—
—
—
—
—
0
Kyushu
23.5
—
22.4
—
0.96
—
1'
Kojima
24.3 ± 0.79
22.5-25.9
24.8 ± 1.09
23.1-26.4
1.02 ± 0.039
0.95-1.10
192
Yakushima
22.1 ± 0.81
20.4-23.3
23.9 ± 1.51
19.1-25.8
1.08 ± 0.065
0.94-1.18
20
Adult males
Honshu
26.1 ± 1.06
23.9-28.2
24.6 ± 1.43
21.7-28.2
0.94 ± 0.054
0.80-1.06
60
Shodoshima
25.0 ± 0.62
24.5-26.2
23.8 ± 0.61
23.2-25.1
0.95 ± 0.031
0.90-1.00
11'
Kyushu
28.2 ± 0.78
27.2-29.2
25.6 ± 0.57
25.0-26.5
0.91 ± 0.042
0.86-0.97
53
Kojima
26.1 ± 1.09
24.2-27.8
25.7 ± 1.30
22.2-27.5
0.98 ± 0.036
0.91-1.03
142
Yakushima
23.6 ± 0.93
21.9-25.3
25.0 ± 1.68
22.6-29.0
1.06 ± 0.066
0.96-1.20
25
' Provisioned population.
^ Semiprovisioned population (ca. 12 kg of wheat grain distributed weekly to ca. 70 monkeys).
^ Includes four provisioned specimens.
samples of 1 1 species of macaques, including two
samples of M. fuscata captives of unknown geo-
graphic origin. Both M. fuscata samples share a
single NRAMPl haplotype, which clusters closely
with the haplotypes of M. mulatta, M. cyclopis,
and M. fascicularis samples; at the center of this
fascicularis-group cluster is a Chinese M. mulatta
haplotype, from which the M. fuscata haplotype
differs by one mutation.
Nuclear DNA: Y-Chromosome — Tosi et al.
(2000, p. 138; 2002, p. 168) sequenced 3.1 kb of
Y-chromosome DNA (SRY and TSPY loci) in
three M. fuscata captives of unknown geographic
origin and compared these sequences with those
in other species of macaques and other catar-
rhines. Y-chromosome DNA in M. fuscata was
found to be generally more similar to that in M.
mulatta and M. cyclopis than to that in M. fasci-
cularis. However, this generalization does not ap-
ply to Indochinese populations of M. fascicularis,
which are postulated to have received M. mulatta
Y-chromosome DNA as a result of hybridization
between dispersing M. mulatta males and philo-
patric M. fascicularis females.
Nuclear DNA: Random Amplified Polymor-
phism— Vernesi et al. (2000, p. 186) have studied
Random Amplified Polymorphic DNA (RAPD) in
samples from nine M. fuscata specimens of un-
known geographic origin and compared these
samples with those from specimens of nine other
macaque species. Within each of the 10 species
studied, including M. fuscata, the RAPD pattern
was identical in all samples. Interspecific cluster
analysis of RAPD patterns suggests that M. fus-
cata is the sister group of a hitherto unrecognized
species group that includes M. nemestrina, M. fas-
cicularis, M. tonkeana, M. arctoides, and M. mu-
latta. However, Vernesi et al. (2000, p. 191) con-
clude their report by cautioning that the uncertain
genetic significance of RAPD variation dictates
that "[p]rudence should guide inferences about
nucleotide divergence, population structure and
phylogeny based on RAPD markers."
Mitochondrial DNA-Nuclear DNA: Multi-
ple Loci— Tosi et al. (2003, p. 1421) have se-
quenced one mitochondrial locus and three nucle-
ar loci in three specimens of M. fuscata (precise
origins unspecified) and 60 specimens of 18 other
macaque species (Table 26). For three of the four
loci (one Y-chromosomal, one mitochondrial, one
autosomal), the sister group o^ M. fuscata was de-
termined to consist variously of one or more fas-
cicularis-group macaque species; for the fourth
locus (autosomal), the sister group of M. fuscata
was determined to consist of four sinica-group
species. Tosi et al. (2003, p. 1425) suggest that
the variation in sister groups that is indicated by
these four loci probably is a result either of in-
complete lineage sorting or of hybridization; for
further discussion of incongruence in molecular
phylogenies that are based on different genes, see
Rokas et al. (2003, p. 798). Based on Y-chromo-
some and mtDNA haplotype data, the divergence
date of M. fuscata, M. mulatta, and M. cyclopis
is estimated to be 1.0-1.2 million years ago (Tosi
et al., 2003, p. 1431).
FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES
35
\ —
D
x:
w
c
O
X
•
A Kojima
□ Kyushu
o Shodoshima
A Yakushima
•
• •
' 4
<
•
< <
<
<<
•
I
males
<
1 o^
<
<
<
Adult
!] ^
o
CO
00
CNJ
CD
CN
CSJ
1
t
#
* 4
•
••
•
smales
# ^
'A
1. %• • i
I*
•
Adult f<
<
^ <
<
< <
< <
•
<
o
o
CO
CO
CM
CD
CN
CNJ
CNJ
O
CNI
s ^
o
■o
1)
eg
ra
o
o
CO
E
4-*
SI
c
k_
O
c
CO
C3
<U
CM
1
>
CO
^
CM
o
Tt"
CSJ
i
(uiuj) ^MBjaq |e)!qjo
36
FIELDIANA: ZOOLOGY
Table 22. Greatest length of skull and rostral-postrostral ratio in fascicularis-gmup species. (References: Table
17; Fooden, 1995, p. 38; 2000, p. 44; Fooden & Wu, 2001, p. 13).
Species
Adult females
Adult males
Mean ± SD
Extremes
N
Mean ± SD
Extremes
Greatest length of skull (mm)
M. fascicularis 100.4 + 5.63 84.0-119.2 439 118.7
M. mulatta 107.8 ± 7.07 92.9-126.5 120 121.8
M. cvclopis 110.4 ±2.40 107.7-114.5 12 125.2
M.fiiscata 119.1 ±4.94 109.2-133.7 141 133.2
Rostral-postrostral ratio (%)
M. fascicularis 47.6 ± 4.21 36.9-61.0 241 56.7
M. mulatta 44.1 ± 3.68 37.6-59.4 111 50.5
M. cyclopis 48.2 ± 3.04 44.7-51.1 4 55.2
M.fiiscata 51.2 ± 3.03 43.5-57.5 137 57.8
7.29
97.4-140.1
454
8.28
107.1-143.1
80
5.20
114.6-133.4
15
5.72
120.6-148.3
100
3.94
41.9-66.3
316
3.04
41.3-58.2
69
2.10
52.6-58.2
8
3.65
48.3-66.2
100
Blood Proteins
The most comprehensive survey of blood-pro-
tein variation in M. fuscata has been pubHshed by
Nozawa et al. (1991, p. 415; 1996, p. 22), who
studied 32 genetic loci in 3409 blood specimens
that originated at 38 localities, spanning the entire
specific range from Shimokita Peninsula to Yak-
ushima (Fig. 20); in addition, these authors com-
pared allele frequencies in M. fuscata with cor-
responding frequencies in a sample of southern
Chinese M. mulatta (n = 150), selected as a close
phylogenetic relative of M. fuscata. A more lim-
ited review of blood-protein variation in M. fus-
cata and M. mulatta was published by Fooden and
Lanyon (1989, p. 215).
Although blood-protein variability apparently is
less in M. fuscata than in many other macaques
(Nozawa et al., 1996, p. 7), intraspecific analysis
of Nei's standard genetic distance (D) has re-
vealed a clear pattern of geographic differentia-
tion in the Japanese species (Fig. 20). Based on
this analysis, Nozawa et al. (1991, p. 425; 1996,
p. 27) allocated the 38 sample localities of M.
fuscata to seven groups; within each of these
groups, interlocality D is less than 0.005.
The geographic distribution of group 1 , which
comprises 28 of the 38 localities, extends broadly
from north-central Honshu to southernmost Kyu-
shu, including the neighboring large island Shi-
koku and small islands Awajishima and Kojima.
The remaining six groups, to which 10 localities
are allocated, are geographically peripheral to
group 1 and are progressively more genetically
distant from it.
Group 2 comprises only one locality (Hakusan)
in central Honshu, slightly marginal to group 1
localities (Fig. 20); between group 1 and group 2,
D ~ 0.01 1. Group 3 comprises three localities on
Izu Peninsula, south-central Honshu; between
group 3 and the two preceding groups, D ~ 0.013.
Group 4 comprises two localities, 220 km apart,
on two small islands (Shodoshima and Kashima)
off the coast of Shikoku; between this group and
groups 1-3, D ~ 0.016. Blood proteins in group
5 — one locality on Boso Peninsula, south-central
Honshu — are somewhat similar (D ~ 0.008) to
those in group 6 — two localities on Yakushima,
ca. 1000 km southwest of Boso Peninsula; be-
tween groups 1-4 and groups 5-6, D ~ 0.025.
Group 7, finally, includes only one locality on
Shimokita Peninsula, northernmost Honshu; be-
tween groups 1-6 and group 7, D ~ 0.035. Al-
though localities in groups 3-7 are restricted to
peninsulas or small islands, this pattern of distri-
bution does not entirely distinguish them from lo-
calities in group 1, some of which are similarly
distributed on peninsulas and small islands (Fig.
20).
Of the 32 blood-protein loci studied by Nozawa
et al., eight are monomorphic and 24 are poly-
morphic, either in M. fuscata or in the Chinese
M. mulatta sample or in both (Table 27). At 19
of the 24 polymorphic loci, the seven M. fuscata
groups are homogeneous with respect to their ma-
jor alleles, and at 15 of these 19 loci, the M. mu-
latta sample also shares the same major alleles.
The four loci at which the common major alleles
in M. fuscata groups differ from the major alleles
in the M. mulatta sample are ADA, Dia, TBPA,
and Tf; high frequencies of the major alleles at
these four loci may be shared derived characters
in M. fuscata.
At 5 of the 24 polymorphic loci, the seven M.
FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES
37
Fig. 18. Geographic variation in mtDNA sequence haplotypes in Macaca fuscata (Hayasaka et al., 1991, p. 401),
sample size in parentheses; for details, see Table 23.
fuscata groups are heterogeneous with respect to
their major alleles, and at all five of these loci,
the major alleles in group 1 are the same as the
major alleles in the M. mulatto sample (Table 27);
this suggests that high frequencies of these alleles
may represent the primitive character state in M.
fuscata. If so, the primary blood-protein diver-
gence of group 7 (the most genetically divergent
of the groups) from the ancestral M. fuscata stock
was an increase in the frequency of allele Hb-(3:
2 (and probably also of allele PGM-II:2), the pri-
mary divergence of groups 5 and 6 was an in-
crease in the frequency of allele PGM-II:2, the
primary divergence of group 4 was an increase in
the frequency of allele CA-I;J„ the primary di-
vergence of group 3 was an increase in the fre-
quency of allele Gc:b, and the primary divergence
of group 2 was an increase in the frequency of
38
FIELDIANA: ZOOLOGY
Table 23.
cf. Fig. 18.
Local variation in mtDNA sequence haplotypes in Macaca fuscata (Hayasaka et al., 1991, p. 401);
Approx.
Reported mtDNA haplotypes
distance
MtDNA
from
]
Interrelated mtDNA
haplotypes
pre-
haplotypes
of uncer-
Locality
(listed in northeast-
ceding
locality
Repeatable sequence status
tain
- relation-
Island
Prefecture
to-southwest order)
(km)
Single Double Triple
ships
N
Group II/IA'II
Honshu
Aomori
Shimokita Peninsula
—
II
1
Honshu
Tochigi
Nikko
510
11(7)'
VI (1)'
8
Honshu
Nagano
Shiga
100
II
15
Honshu
Kanagawa
Izu Peninsula
230
V
1
Honshu
Fukui
Takahama
320
P VIP
18
Honshu
Kyoto
Arashiyama
60
II
6
Honshu
Osaka
Minoo-city
20
II
1
Honshu
Tottori
Wakasa
120
II
8
Shodoshima
Kagawa
Shodoshima
90
II
Group III/IVA^III
2
Honshu
Okayama
Gagyusan
60
111
5
Kyushu
Oita
Takasakiyama
250
V
3
Yakushima
Kagoshima
Yakushima
340
IV3 VHP
32
' Dimorphic.
' Heteroplasmic (12/18).
^ Heteroplasmic (29/32).
allele MDH:2. Changes in frequency of minor al-
leles apparently have played a lesser role in the
genetic differentiation of M. fuscata groups (No-
zawa et al., 1991, p. 427; 1996, p. 27).
Independent random genetic drift in semi-iso-
lated local populations probably accounts for
much of the observed blood-protein variation in
M. fuscata (Nozawa et al., 1991, p. 431; 1996, p.
32). Following are supplementary comments con-
cerning two striking disjunctions in blood-protein
group distribution:
1. Group 4 populations, characterized by rela-
tively high frequency of allele CA-l-d^, apparently
are restricted to Shodoshima and Kashima, two
small islands that are more than 200 km apart
(Fig. 20); between these two islets is Shikoku, a
large island apparently inhabited by group 1 pop-
ulations. Two alternative hypotheses may apply
here: (a) high frequencies of CA-I-ti, evolved in-
dependently on Shodoshima and Kashima, or (b)
CA-I-^2 populations formerly were widespread on
Shodoshima, Shikoku, and Kashima but subse-
quently were replaced on Shikoku by group 1
populations with high frequency of CA-I-a (cf.
Nozawa et al., 1991, p. 431; 1996, p. 31).
2. Groups 5 and 6 populations, characterized
by high frequencies of allele PGM-II:2, are dis-
tributed ca. 1000 km apart on Boso Peninsula
(south-central Honshu) and Yakushima (southern-
most island inhabited by M. fuscata), respectively
(Fig. 20); PGM-II:2 also is relatively frequent
(41.1%) in group 7 populations on Shimokita Pen-
insula (northernmost Honshu). As above, two al-
ternative explanatory hypotheses may apply here:
(a) high frequencies of PGM-II:2 evolved inde-
pendently on Shimokita Peninsula, Boso Penin-
sula, and Yakushima, or (b) PGM-II:2 formerly
was widespread on the Japanese archipelago but
subsequently has virtually disappeared every-
where except on Shimokita Peninsula, Boso Pen-
insula, and Yakushima (cf. Hayasaka et al., 1987,
p. 513; Nozawa et al., 1991, p. 433; 1996, p. 32).
The pattern of geographic variation of blood-
protein groups in M. fuscata (Fig. 20) differs
strikingly from the pattern of geographic variation
of mtDNA types in this species (Figs. 18-20).
Three major differences may be noted: (1) unlike
blood-protein groups, mtDNA types are abruptly
discontinuous in west-central Honshu; (2) unlike
blood-protein groups, mtDNA type on Shimokita
Peninsula is not strongly divergent from that else-
where on Honshu; and (3) also unlike blood-pro-
tein groups, mtDNA type on Shodoshima is not
divergent from that in neighboring central Hon-
FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES
39
8
2
9
4
10
11
35
36
37
38
3
38 + 1
1
39
2
40
2
41
2
41 - 1
2
41 + 1
1
42
1
43
1
Table 24. Mitochondrial DNA haplotypes detected in Macaca fiiscata at localities sampled by Kawamoto (2002,
p. 60); cf. Fig. 19.
Haplotype
no. N Localities'
Group I haplotypes
Kurokawa-mura
Arakawa-mura; Yokoze-machi
Ashigakubo, Hinohara-mura; Itsukaichi; Tabayama-mura 1
Okutama-machi
Tabayama-mura 2
Awajishima
Gagyusan
Anan-city
Hiromi-cho; Nakamura-city; Nametoko
Matsuno-cho
Kisuki-cho; Ooda-city
Kugino-mura; Takasakiyama
Kanoya-city; Kojima
Sagara-mura; Satsuma-cho
Kushima-city
Yakushima 1
Yakushima 2
Group II haplotypes
21 Ajigasawa-machi; Asahi-mura; Fukushima-city; Genbikei; Goyosan 1; Hachimori-machi;
Higashine-city; Kaminoyama-city; Kanita-machi; Kashima-machi; Kinkazan; Ogachi-ma-
chi; Shichigashuku-machi; Shimokita 1; Shimokita 2; Shimokita 3; Shimokita 4; Soma-
city; Takahatamachi; Tsugaru-touge; Yamadera
Goyosan 2
Nikko 1
Nikko 2
Boso
Kyonan-machi
Tsukui-machi
Nishiizu-cho
Atami-city; Odawara-city
Aikawa-machi; Fujiyoshida-city; Kiyokawa-mura 1; Nishikatsura-cho
Sanada-machi; Shiga-kogen; Sugadaira
Karuizawa-machi
Hodaka 1 ; Omachi-city; Outaki-mura
Kamiichi-machi
Oosawano-machi
Hodaka 2
Unazuki-machi 1
Arashiyama 1; Asahi; Hosoiri-mura; Mihama-cho, Mikata-gun; Nishiazai-cho; Sodoshima;
Unazuki-machi 2
Neo-mura
Inuyama-city; Kiyokawa-mura 2; Tatsuyama-mura; Shinshiro-city; Toei-cho
Mukawa-mura
Shichisou-cho
Takahama-cho
Adogawa-cho 3^
Hatasho-cho
Eigenji-cho; Hino-cho; Kameyama-city; Tsuchiyama-cho
Hokusei-cho; Inabe-cho; Taga-cho
Kozagawa-cho
Hongu-cho
Kanaya-cho; Koka-cho 1 ; Kumano-city; Mihama-cho, Minamimuro-gun; Owase-city; Oya-
mada-mura; Tsubaki
Adogawa-cho 1 ; Adogawa-cho 2; Arashiyama 2; Hieizan; Ibuki-cho; Kohoku-cho; Koka-
cho 2; Minoo-city; Natasho-mura; Otsu-city; Sasayama-cho; Takatsuki-cho; Yogo-cho
Wakasa-cho
' For details, see Gazetteer, Appendix 4; two specimens were sampled at each of nine localities (indicated by number
"1" or "2" following a locality name), three specimens were sampled at one locality (Adogawa-cho 1-3), and four
specimens were sampled at one locality (Shimokita 1-4).
- Y. Kawamoto (pers. comm., Nov. 2003).
40 FIELDIANA: ZOOLOGY
2
1
3
1
4
1
5
1
6
1
12
1
13
1
14
2
15
4
16
3
16 + 1
1
17
3
17 + 1
1
17 + 2
1
18
1
19
1
20
7
20 + 1
1
21
5
22
1
23
1
26
1
26 + 1
1
27
1
28
4
29
3
30
1
31
1
32
7
33
13
34
1
130°
135'
Fig. 19. Geographic variation in mtDNA sequence haplotypes in Macaca fuscata (Kawamoto, 2002, p. 60); for
details, see Table 24.
shu. The discrepancies between the distributions
of these two types of genetic characters presum-
ably are related to their respective modes of in-
heritance— biparental for blood proteins and uni-
parental (maternal) for mtDNA types (cf. Melnick
et al., 1993, p. 290; Tosi et al., 2000, p. 138).
Because female macaques are much more philo-
patric than males, mtDNA types would be ex-
pected to disperse more slowly than blood pro-
teins; the distribution of mtDNA types therefore
would be expected to preserve evidence of more
ancient zoogeographic barriers or dispersal routes.
Except for the distinctiveness of the Yakushima
population, neither blood-protein group distribu-
FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES
41
Table 25. Mean mtDNA divergence estimates reported between Macaca fuscata and M. mulatta, M. cyclopis,
and M. fascicularis.
Mean divergence (Div.) estimates (%)
References
Analytical
techniques
Intra-
specific
M. fuscata M. mulatta
Divergence from M. fuscata
M.
cyclopis
M. fasci-
cularis
Div. N Div. N Div. N Div. N
Hayasaka et al. (1988b. p. 277)
Hayasaka et al. (1988a. p. 635)
Hayasaka et al. (1996, p. 1049)
Zhang & Shi (1989, p. 335)
Zhang & Shi (1993. p. 12)
Melnick et al. (1993, p. 286)
Morales & Melnick (1998, p. 13)^
Kawamoto et al. (1999. p. 57)
LP
17 enzymes
1.6
10
3.6
1
3.7
1
6.2
1
uencing: 0.9 kb'
—
1-
3.7
1
—
0
8.8
1
uencing: 0.896 kb'
1.6
2"*
3.6
1
3.2
1
9.4
1
LP
10 enzymes
—
35
2.8
4
—
0
—
0
LP
16 enzymes
—
P
3.8
2
—
0
6.5
1
LP
15 enzymes
1.7
10^
3.7
19"
3.9
V
6.0
P
LP
22 enzymes
0
2
0.6
3
1.0
2
3.2
3
ue
ncing: 0.464 kb*
—
1
—
0
2.4
1
—
0
' Includes genes for tRNA"'\ tRNA^", tRNA^*^", 3' region of ND4, and 5' region of ND5.
- One of 10 specimens previously studied by Hayasaka et al. (1988b, p. 277).
' Hindlll fragment.
■* Includes one of 10 specimens previously studied by Hayasaka et al. (1988b. p. 277).
' Data from Hayasaka et al. (1986, p. 352: 1988b, p. 277).
* Includes data from one specimen reported by Hayasaka et al. (1988b, p. 277).
■^ Cf. Tosi et al. (2002. p. 168). who sequenced a 1.5-kb fragment of mtDNA (3' end of 12S ribosomal gene;
tRNA^'^'; 5' end of 16S ribosomal gene) in M. fuscata (n = 3), M. mulatta (n = 5), M. cyclopis (n = 2), and M.
fascicularis (n - 17); although divergence estimates are not specified in this work, the authors' bootstrap consensus
tree (fig. 2a) indicates that mtDNA in M. fuscata is more similar to that in M. mulatta and M. cyclopis than to that
in M. fascicularis.
^ D loop.
tions nor mtDNA type distributions bear much re-
lationship to the present configuration of land and
water in the Japanese archipelago (Figs. 18-20).
Instead, as indicated above, these distributions
may reflect the climate, vegetation, land bridges,
and water gaps that existed during previous stages
in the evolutionary history of M. fuscata.
Hemoglobin Amino-Acid Sequence
The amino-acid sequence of hemoglobin in M.
fuscata has been compared with that in M. mu-
latta (Matsuda, 1982, p. 293). In these species,
the a-chains of hemoglobin are identical, and the
P-chains differ only in a single amino acid.
Table 26. Sister-group relationships of Macaca fuscata indicated by sequencing of paternal, maternal, and bi-
parental markers (Tosi et al.. 2003. pp. 1426-1429).
Loci studied
M. fuscata
specimens'
Indicated sister group-
'-chromosome:
Nos. 29-
-31
SRY, TSPY, 3. 1 kb
ItDNA:
Nos. 29-
-31
12s ribosomal gene, 1.5 kb
kUtosome:
Nos. 29-
-31
C4 long intron 9, 3.3 kb
LUto.some:
No. 29'
IRBP intron 3, 1.6 kb
No. 31'
(CYC (MUL India (MUL Burma/China, FAS Thailand/
Cambodia/Vietnam)))
(MUL Burma)
((SIN)(RAD)(ASS. THD)
(FAS West Malaysia)
(FAS Thailand)
' Origin "Japan," precise localities unspecified.
- Key to species abbreviations: /a.vc/cM/am group — FAS = M. fascicularis. CYC = M. cyclopis. MUL = M. mulatta:
sinica group — SIN - M. sinica, RAD = M. radiata. ASS = M. assamensis, THI = M. thibetana.
-' Sister-group relationships of M. fuscata specimen no. 30 are not reported for this locus.
42
FIELDIANA: ZOOLOGY
130°
135°
140°
Fig. 20. Geographic distribution and genetic interrelationships of blood-protein groups in Macaca fuscata (Nozawa
et al., 1991, p. 418; 1996, p. 4); for details, see Table 27.
Antibodies to Cedar Pollen Antigens
Natural allergy to the pollen of Japanese cedar
(Cryptomeria japonica) has been detected in M.
fuscata (Nakamura et al., 1991, p. 643). In six
provisioned free-ranging groups and three captive
groups studied in nine prefectures (n = 276), the
incidence of antibodies to crude cedar pollen an-
tigen averaged 16.3% and varied from 0% to
31.3% (Table 28). The 45 positive sera were fur-
ther tested for reactivity to two purified cedar pol-
len antigens. Cry j I and Cry j II; of the positive
sera, 23 reacted to both antigens, 21 reacted only
to Cry j I, and one reacted only to Cry j II. The
factor or factors responsible for prefectural vari-
ation in M. fuscata reactivity to cedar pollen an-
FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES
43
Table 27. Mean frequency' (%) of major alleles at 24 loci in blood-protein groups of Macaca fuscata (cf. Fig.
20) and frequency of corresponding alleles in southern Chinese M. mulatta sample;^ for details, see Nozawa et al.
(1991, p. 418; 1996, p. 4).
Fre-
Blood-protein groups^ and mean frequencie
«
quency
(%) in
M.
-
of major alleles in M. fuscata
Group
mulatta
Group 2:
Group
7: Shi-
sample:
Group 1:
central
Group 3:
Group 4:
5: Boso
mokita southern
widely
Honshu
Izu
Kashima and
Penin-
Group 6:
Penin-
China
Major
distributed
(1 local-
Peninsula
Shodoshima
sula (1
Yakushima
sula (1
(n =
Locus
allele
(28 localities)
ity)
(3 localities)
(2 localities)
locality) (2 localities) locality)
150)
Loci at which M.
. fuscata \
groups are heterogeneous with
I respect
to major alleles
CA-I
a
97.0 ± 4.5
85.9-100.0
100.0
100.0
34.6
17.6-51.6
100.0
100.0
100.0
93.0
di
2.3 ± 4.0
0-14.1
0
0
65.4
48.4-82.4
0
0
0
5.3
Gc
a
95.6 ± 5.8
78.6-100.0
76.9
45.7 ± 13.9
35.7-61.5
100.0
100.0
100.0
78.8
98.0
h
2.1 ± 5.3
0-21.4
1.9
54.1 ± 14.3
37.7-64.3
0
0
0
21.2
0
Hb-3
1
99.9 ± 0.4
98.4-100.0
100.0
85.6 ± 8.6
78.5-95.1
100.0
100.0
100.0
14.3
100.0
2
0.1 ± 0.4
0-1.6
0
14.4 ± 8.6
4.9-21.5
0
0
0
85.7
0
MDH
1
97.3 ± 4.5
84.4-100.0
48.1
95.1 ± 3.5
91.8-98.7
100.0
100.0
100.0
91.4
100.0
2
1.5 ± 3.4
0-14.3
51.9
3.9 ± 3.7
1 .3-8.2
0
0
0
0
0
PGM-II
1
99.7 ± 0.9
96.4-100.0
100.0
100.0
97.7
95.4-100.0
44.2
8.3
2.6-13.9
52.9
96.3
2
0.3 ± 0.9
0-3.6
0
0
0
55.8
91.7
86.1-97.4
47.1
3.7
Loci at which M
. fuscata :
groups are homogeneous with
respect to major alleles
Acp
A
99.1 ± 2.3
89.3-100.0
100.0
100.0
100.0
100.0
100.0
100.0
98.7
ADA^
I
100.0
100.0
100.0
100.0
100.0
100.0
100.0
0
AK
I
98.1 ± 3.8
85.0-100.0
100.0
98.6 ± 1.3
97.4-100.0
99.4
98.9-100.0
100.0
100.0
100.0
99.0
Alb
A
100.0 ± 0.01
99.9-100.0
88.5
100.0
100.0
100.0
100.0
100.0
97.7
Alp
A
100.0
100.0
100.0
100.0
99.5
100.0
100.0
100.0
CA-II
a
99.2 ± 2.2
90.5-100.0
82.7
100.0
100.0
71.8
100.0
85.7
97.3
Cell Es
1
98.4 ± 3.5
87.5-100.0
100.0
83.1 ± 10.4
76.1-95.1
100.0
100.0
100.0
100.0
99.3
ChEs
I
99.9 ± 0.2
99.2-100.0
100.0
100.0
99.9
99.8-100.0
100.0
100.0
100.0
100.0
Dia^
A
99.8 ± 1.0
94.6-100.0
100.0
100.0
100.0
100.0
100.0
100.0
13.4
Es-D
1
99.7 ± 1.2
95.0-100.0
100.0
98.4 ± 2.8
95.1-100.0
100.0
100.0
100.0
100.0
99.7
IDH
2
100.0 ± 0.01
99.9-100.0
100.0
100.0
100.0
100.0
100.0
100.0
82.9
LDH-A
1
98.1 ± 3.6
84.2-100.0
96.2
100.0
100.0
100.0
100.0
100.0
100.0
LDH-B
I
99.9 ± 0.3
98.2-100.0
100.0
100.0
93.4
86.8-100.0
100.0
100.0
100.0
100.0
44
FIELDIANA: ZOOLOGY
Table 27. Continued.
Fre-
Blood
l-protein groups' and mean frequencies
quency
(%) In
M.
-
of major alleles in M. fuscata
Group
mulatto
Group 2:
Group
7: Shi-
sample:
Group 1:
central
Group 3:
Group 4:
5: Boso
mokita southern
widely
Honshu
Izu
Kashima and
Penin-
Group 6:
Penin-
China
Major
distributed
(1 local-
Peninsula
Shodoshima
sula (1
Yakushima
sula (1
(n =
Locus
allele
(28 localities)
ity)
(3 localities)
(2 localities)
locality) (2 localities) locality)
150)
PGM-1
1
99.8 ± 0.8
95.8-100.0
100.0
91.5 ± 4.1
86.9-94.9
100.0
100.0
100.0
100.0
98.7
PHI
1
94.2 ± 7.9
73.8-100.0
100.0
100.0
99.9
99.7-100.0
81.4
100.0
82.9
97.3
Pi
C
97.7 ± 4.7
80.4-100.0
100.0
100.0
99.7
99.5-100.0
94.2
100.0
100.0
98.7
6PGD
A
100.0
100.0
100.0
100.0
100.0
100.0
100.0
97.6
TBPA
S
100.0
100.0
100.0
100.0
100.0
100.0
100.0
25.0
TP
F
96.3 ± 5.3
77.9-100.0
92.3
81.3 ± 4.9
78.3-86.9
100.0
100.0
100.0
100.0
8.7
' Unweighted mean of local allele frequencies within each M. fuscata group; standard deviation and extremes also
specified, where pertinent.
- Excludes eight loci monomorphic in M. fuscata and M. mulatta samples: a,; Amy; Cat; Cp; G6PD; Hb-a; Hp;
To.
' Group localities (island name in capitals, prefecture name in italics, locality name in parentheses, sample size in
square brackets): Group 1 [2566 samples]— AWAJISHIMA, Hyogo (Awajishima [150]); HONSHU, Aichi (Oshiro
[42]), Fukui (Mikata [59], Takahama [49]), Fukushima (Fukushima-city [31]), Hiroshima (Kochi [34]), Hyogo (Ka-
suga-cho [70]), Kyoto (Arashiyama [228], Hiyoshi-cho [49], Ine-cho [30], Wachi-cho [31]), Nagano (Shigakogen
[208]), Nara (Muroji [19]), Okaxama (Gagyusan [172], Katsuyama-cho [28]), Osaka (Minoo-city [19]), Shiga (Ryo-
zenyama [120]), Shizuoka (Tenryu-city [12]), Tochigi (Nikko [21]), Tottori (Wakasa [25]); KOJIMA, Miyazaki (Koji-
ma [88]); KYUSHU, Fukuoka (Kawaradake [52]), Kumamoto (Itsuki-mura [21]), Miyazaki (Kushima-city [28]), Oita
(Kamae-machi [30], Takasakiyama [866]); SHIKOKU, Ehime (Nametoko [26]), Kochi (Odo [58]). Group 2 [26
samples] — HONSHU, Ishikawwa (Hakusan [26]). Group 3 [297 samples] — HONSHU, Kanagawa (Yugawara-machi
T [61]), Shizuoka (Hagachi [158], Ihama [78]). Group 4 [215 samples]— KASHIMA, Ehime (Kashima [17]); SHO-
DOSHIMA, Kagawa (Shodoshima [198]). Group 5 [94 samples]— HONSHU, Chiha (Takagoyama [94]). Group 6
[176 samples]— YAKUSHIMA, Kagoshima (Kamiyaku-cho [54], Yaku [122]). Group 7 [35 samples]— HONSHU,
Aomori (Wakinosawa [35]). Total samples, 3409.
^ Allele frequencies at these three loci recently have been studied in 20 blood samples collected at four localities
in Mie Prefecture (HONSHU: Aoyama; Inabe-cho; Kiwa; Omiya) and four localities in Wakayama Prefecture (HON-
SHU: Nakatsu; Kanaya-cho; Kozagawa-cho; Motomia) (Kawamoto et al., 1999, p. 55; 2001, p. 15); reported fre-
quencies are: ADA, allele /—100.07c; Dia, allele A— 100.0%; and Tf, allele F— 92.5%, allele G— 7.5%.
tigens have not been determined (Hashimoto et
al., 1994, p. 396).
Karyology
Classical staining techniques revealed that the
diploid chromosome number in M. fuscata is 42
and that chromosome morphology in this species
is generally similar to that in other macaques and
also to that in other papionins (Chiarelli, 1962, p.
410). Although banding techniques subsequently
confirmed these general similarities, one species-
specific pericentric inversion was discovered on
chromosome 15 oi M. fuscata (n == 10) (Stanyon
et al., 1983, p. 142; 1988, p. 6); banding tech-
niques also revealed that M. fuscata, like some
other papionin species, is polymorphic with re-
spect to size of the nucleolar organizer region on
chromosome 1 3 and size of the C band on the Y
chromosome.
In a survey to determine whether congenital
limb malformation in M. fuscata is related to
chromosomal abnormality, Minezawa et al. (1990,
p. 572) studied the karyotypes of 257 monkeys
representing seven local populations on five Jap-
anese islands (Awajishima, n = 28; Honshu, n =
95, three populations; Kojima, n = 60; Kyushu,
n = 53; Shodoshima, n = 21). No relationship
was found between limb malformation and chro-
FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES
45
Table 28. Incidence of serum specimens positive for antibodies to crude cedar pollen antigen and incidence of
serum specimens with positive reactions to purified cedar pollen antigens (Cry j I, Cry j II) in prefectural samples of
Macaca fuscata. '
Positive for
antibodies to
crude antigen
Positive reactions to
1 purified antigens
Prefecture
Cryj I and Cry j II
Cryj
I only
Cryj II only
N
Honshu
Nagano
Gunma
Tokyo
Aichi
Kyoto
Okayama
Osaka
2(9.1%)
8 (27.6%)
1 (5.0%)
5(16.1%)
5(31.3%)
0
I (6.3%)
0
5
1
2
1
0
0
2
2
0
3
4
0
1
0
1
0
0
0
0
0
22
29
20
31
16
25
16
Awajishima
Hyogo
13(25.5%)
10
Kojinia
3
0
51
Miyazaki
10(15.1%)
4
6
0
66
Totals
45(16.3%)
23(51.1%)
21 (46.7%)
1 (2.2%)
276
Reference: Hashimoto et al. (1994, p. 395; cf. Nakamura et al.. 1991. p. 645: Sakaguchi et al.. 1992. p. 325).
mosomal abnormality (cf. Iwamoto & Hirai,
1970, p. 396), and no geographic variation in kar-
yotype was reported.
Parasites
Protistans
One species of flagellate and four species of
amebas have been detected at an incidence of 20-
75% in fecal samples of M. fuscata at Shimokita
Peninsula, which is at the northern limit of the
monkey's distribution (Table 29). The ciliate Bal-
antidium coli has been recorded in a M. fuscata
captive by Tschemer (1985, p. 226).
Malarial parasites (sporozoans) apparently are
naturally absent from M. fuscata, judging from
547 blood specimens collected on Honshu, Sho-
doshima, Miyajima, Kyushu, and Yakushima (O-
tsuru & Sekikawa, 1979, p. 347). The absence of
malaria in M. fuscata presumably is a conse-
quence of Japan's geographic location, which is
north of the range of the Leucosphyrus group of
anopheline mosquitoes; this group of mosquitoes
apparently is essential in the transmission of sim-
ian malaria in Asia (Coatney et al., 1971; Fooden.
1994, p. 582). M. fuscata is susceptible to exper-
imental infection with Plasmodium coatneyi (Ka-
wai et al., 1998, p. 101), which is a natural ma-
larial parasite of M. fascicularis in Southeast Asia
(Fooden, 1994, p. 576).
The sporozoan Babesia sp. has been detected
in 4 of 93 blood specimens (4.3%) collected from
members of a M. fuscata group that had been
translocated in 1957 from Yakushima to Honshu
(Ohirayama) (Otsuru & Sekikawa, 1979, p. 347).
This sporozoan has not been detected in nontran-
slocated groups of M. fuscata, either on Yakushi-
ma (n = 39) or on Honshu, Shodoshima, Miya-
jima, and Kyushu (n = 415).
Table 29. Incidence of protistan parasite species de-
tected in fecal samples of Macaca fuscata on Shimokita
Peninsula, Aomori Prefecture (n = 20).'
Incidence
Species
No.
%
Flagellate
Giardia sp.
10.0
Amoebas-
Eiuamoeba coli 5
25.0
E. histolytica 15
75.0
Endolimax nana 14
70.0
lodamoeba buetschlii 5
25.0
' Reference: Takada et al. (1981, p. 70).
- The same four species of amoebas have been detect-
ed in fecal samples of a laboratory colony of M. fuscata
(Koyama et al.. 1978. p. 60).
46
FIELDIANA: ZOOLOGY
Table 30. Incidence of cestode parasites (Bertiella studeri, possibly also other species) in local samples' of
Macaca fuscata.
Cestode
Latitude
incidence
Locality
Island
(N)
(%)
N
References^
Shimokita'
Honshu
41°10'
0
35
1,2
Kinkazan
Kinkasan
38°17'
0
23
2
Nikko
Honshu
36°44'
0
23
2
Shiga
Honshu
36°44'
0
141
2,3,4
Hakusan"*
Honshu
36°15'
0
25
2
Mikata
Honshu
35°33'
3.2
63
4
Kiso
Honshu
35°31'
0
7
2
Takagoyama
Honshu
35°13'
0
24
2
Sakamoto
Honshu
35°08'
12.5
8
2
Arashiyama
Honshu
35°00'
20.5
156
2,4,5
Hasumi
Honshu
34°53'
0
47
4
Hagachizaki
Honshu
34°4r
0
38
2
Shodoshima
Shodoshima
34°30'
0
95
2,4
Kochi
Honshu
34°28'
0
34
4
Mihara-city
Honshu
34°24'
0
119
4
Awajishima
Awajishima
34°14'
0
20
2
Takasakiyama
Kyushu
33°10'
42.6
155
4,6
Misho/Nishiumi
Shikoku
32°58'
0
36
7
Kojima
Kojima
31°27'
>8.05
286
2, 4, 8, 9
Yakushima''
Yakushima
30°30'
3.8
157
2,4
Totals
—
—
-8.7
1492
—
' Fecal samples, except Sakamoto and Misho/Nishiumi, which are necropsy samples.
2 Key to references: /. Takada et al. (1981, p. 70). 2. Gotoh (2000, p. 294). 3. Nigi et al. (1975, p. 36). 4. Nigi
(1983, p. 38). 5. Machida and Sano (1974, p. 77). 6. Kagei and Hasegawa (1974, p. 236). 7. Itoh et al. (1988, p.
236). 8. Kawai (1974, p. 17). 9. Horii et al. (1982, pp. 416, 418).
^ Includes Wakinosawa.
^ Includes Kamuri.
' Cestode eggs reported in 23 of 73 samples (Reference 8), and "few cestode segments" reported in 74 samples
(reference 9); 139 samples negative for cestodes.
^ Cf. Tanaka et al. (1962, p. 115) and Hayama and Nigi (1963, p. 103), who report B. studeri in two of 66 Yakushima
captives at the Japan Monkey Centre, Inuyama (Honshu), and Nigi (1982, p. 248), who reports no cestodes in 57
Yakushima monkeys from a population that had been translocated to Ohirayama (Honshu).
Trematodes
No flukes were detected in parasitological nec-
ropsy of 36 M. fuscata specimens collected at
Misho/Nishiumi, Ehime Prefecture, Shikoku (Itoh
et al., 1988, p. 236).
Cestodes
north of ca. 36°N. The incidence of tapeworm par-
asites in necropsy samples (1/44 = 2.3%) appar-
ently is somewhat less than that in fecal samples
(129/1448 = 8.9%).
Tscherner (1985, p. 226) has reported that a M.
fuscata captive was infected with the tapeworm
Hymenolepis diminuta, normally a parasite of ro-
dents.
Tapeworm infections are relatively rare — 8.7%
incidence — in natural populations of M. fuscata
(Table 30), and only one species of tapeworm,
Bertiella studeri, has been positively identified as
a natural parasite in these monkeys. The geo-
graphic distribution of tapeworms in M. fuscata
generally appears sporadic, although it may be
significant that no infection has been recorded in
the cool, northern part of the monkey's range.
Nematodes
Six species of nematodes are known to parasit-
ize natural populations of M. fuscata (Table 31);
this excludes two nematodes {Ternidens deminu-
tus and Oxyuridae gen. sp.) reported in captive
M. fuscata by Tscherner (1985, p. 227). Of the
nematode parasites in natural populations, Trich-
uris trichiura is the most prevalent (60.1% inci-
FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES
47
Table 31. Incidence {%) of nematode parasites in local samples' of Mocaca fuscata.
Strep-
Oeso-
Gongylo-
to-
Latitude
Stron-
phago-
nenma
phara-
Locality
Island-
(N)
Trichuris
gyloides
stomiim
spp.
gus'
N
References^
Shimokita"
H
41°10'
60.0
48.6^
0
0
0
35
1.^ 2
Kinkazan
Ki
38°I7'
91.3
13.0
8.7
0
0
23
2
Nikko
H
36°44'
100.0
0
0
0
0
23
2
Shiga
H
36°44'
65.7
0
0
0
0
216
2, i," 4, 5'o
Hakusan"
H
36°15'
63.1
0
23.1
0
1.5
65
2, 4
Mikata
H
35°33'
52.4
15.9
44.4
—
22.2
63
5'o
Kiso
H
35°3r
100.0
71.4
0
0
0
7
2
Takagoyama
H
35° 13'
20.8
16.7
8.3
0
0
24
2
Sakamoto
H
35°08'
87.5
25.0
100.0
0
75.0
8
2
Arashiyama
H
35°00'
75.6
41.7
3.8
0
69.9
156
2. 5:^' 6
Hasumi
H
34°53'
76.6
46.8
0
—
23.4
47
5'o
Hagachizaki
H
34°4r
92.1
42.1
0
0
78.9
38
2
Shodoshima
Sho
34°30'
67.4
16.8
20.0
0
67.4
95
2, 5,'° 7
Kochi
H
34°28'
58.8
38.2
2.9
—
76.5
34
J'o
Mihara-city
H
34°24'
81.5
44.5
5.9
—
42.0
119
5'o
Awajishima
A
34° 14'
95.0
55.0
0
0
20.0
20
2
Takasakiyama
Ky
33° 10'
67.9
30.5
86.2
—
85.6
305
4. 5.'<'S
Misho/Nishiumi
Shi
32°58'
36.1'-
36.1'-
5.6
—
69.4
36
9
Aira
Ky
31°44'
—
—
—
>0"
—
•7
W
Kojima'^
Ko
31°27'
49.6
35.1
31.9
0
87.0
276
2. 4, J.'o 77'5
Toimisaki
Ky
31°22'
0
43.8
69.6
—
72.3
112
4
Yakushima'^
Y
30°30'
45.2
42.0
29.9
10.2
43.9
157'^
2, 5'o
Totals
—
—
60.1
29.9
30.4
<1.0
53.3
1859'^
—
' Fecal samples, except Sakamoto. Misho/Nishiumi. and Aira. which are necropsy samples.
- Abbreviations: A = Awajishima; H = Honshu; Ki = Kinkasan; Ko = Kojima; Ky = Kyushu; Shi = Shikoku;
Sho = Shodoshima; Y = Yakushima.
' Goiigylonema macrogubernacuhim and/or G. pukhrum (see reference 2). Excluded are Gongylonema infections
reported in translocated captives by Uni et al. (1994. p. 128); as indicated by Gotoh (2000. p. 296). some or all of
these infections may have been acquired in captivity.
■* Supplementary record of this species (Yamaguti. 1935. p. 453): "in the small intestine of a Japanese monkey
from Nara Prefecture," 33°5r-34°44'N.
'^Key to references: /. Takada et al. (1981. p. 70). 2. Gotoh (2000, pp. 294. 296. cf. 1989, p. 9). 3. Nigi et al.
(1975. p. 36). 4. Usui and Horii (1982. p. 272). 5. Nigi (1982. p. 248; 1983. p. 38). 6. Machida and Sano (1974. p.
77); Machida et al. (1978. p. 1). 7. Tanaka et al. (1963. p. 93). 7. Kagei and Hasegawa (1974. p. 236). 9. Itoh et al.
(1988. p. 236). 10. Uni et al. (1992, p. 221). 11. Kawai (1974, p. 17).
*' Includes Wakinosawa.
^ Cited in reference / as Strongyloides sp. near fulleborni.
** Absence of O. aculeatum. Gongylonema sp., and S. pigmentotus inferred.
'* In this reference, parasites, except T. irichiiira, are cited by generic names only.
'° In this reference, Trichuris trichiura is cited as Trichocephalus trichiiirus.
" Includes Kamuri.
'- Corrected value; apparent miscalculation in cited reference.
'■' One monkey reported positive for G. pukhrum.
'■*Cf. Horii et al. (1982, pp. 419-421).
'■* In this reference, parasites are cited by generic names only.
"'Cf. Hayama and Nigi (1963. p. 103). Tanaka and Nigi (1967. p. 99). and Usui and Horii (1982. p. 272). who
have investigated the incidence of nematode parasites in captive and translocated populations of Yakushima M.
fuscata.
' '' For Gongylonema spp.. n = 59.
'** For Gongylonema spp., n = 1761.
dence) and is the only nematode parasite that is
widely distributed in northern populations of M.
fuscata; Streptopharagus pigmentatus is next
most prevalent (53.6%). but this species is rare
north of 35°08'N; Oesophagostomum aculeatum
and Strongyloides fuelleborni are relatively un-
common (30.47r and 29.9%. respectively); and
Gongylonema macroguhernaculum and G. pul-
48
FIELDIANA: ZOOLOGY
chrum apparently are restricted to M. fuscata pop-
ulations that inhabit the southern islands Kyushu
and Yakushima (10.2% incidence of Gongyione-
ma spp. in a sample of 59 Yakushima M. fuscata).
Heavy snowfall and low winter temperatures in
the northern part of the range of M. fuscata ap-
parently inhibit intermonkey transmission of most
nematode parasites, except T. trichiura, which is
transmitted by means of a resistant thick-walled
egg (Nigi et al., 1975, p. 45; Usui & Horii, 1982,
p. 273; Horii, 1983, p. 14; Gotoh, 1989, p. 9;
2000, pp. 292, 296).
Although Hayama and Nigi (1963, p. 103) have
tabulated Physaloptera caucasica and Physalop-
tera sp. as nematode parasites of captive Yaku-
shima M. fuscata, this appears to be a typograph-
ical error. In their text (p. 102), these authors
clearly specify that P. caucasica and Physalop-
tera sp. were detected only in M fascicularis and
Papio cynocephalus, not in M. fuscata, which di-
rectly contradicts the corresponding tabular en-
tries (cf. Tanaka & Nigi, 1967, p. 103).
Seasonal variation of nematode parasite inci-
dence in M. fuscata on Kojima has been studied
by Imada et al. (1977, p. 15), Horii et al. (1982,
p. 420), and Nigi (1982, p. 248; 1983, p. 39). On
Kojima and possibly elsewhere, the incidence of
T. trichiura, Streptopharagus pigmentatus, and O.
aculeatum tends to decrease in summer, reaching
a minimum in August, and tends to increase in
winter, reaching a maximum in January-February
(cf. Usui & Horii, 1982, p. 270). Evidence con-
cerning seasonal variation in the incidence of
Strongyloides fuelleborni is equivocal.
The incidence of at least some nematode par-
asites apparently also varies with the age of the
host monkey (Imada et al., 1977, p. 15; Horii et
al., 1982, p. 419; Gotoh, 2000, p. 295). In M.
fuscata on Kojima, for example, the incidence of
Strongyloides fuelleborni in samples of immatures
(age <5 years; combined n = 187) is 75-100%,
whereas the incidence in samples of old adults
(age >13 years; combined n = ca. 120) is ca. 19-
30%.
Sexual variation of the host does not appear to
be an important factor in the incidence of nema-
tode parasites in M. fuscata (Horii et al., 1982, p.
418; Gotoh, 2000, p. 295).
Mite
A "mite egg?" has been reported in one of 20
M. fuscata fecal samples collected on Shimokita
Peninsula, northern Honshu (Takada et al., 1981,
p. 70).
Lice
Pedicinus (Parapedicinus) obtusus japonicus
has been recorded as an ectoparasite of M. fuscata
at Kamikochi (Nagano Prefecture) and "Province
of Shinano" (= Nagano Prefecture) by Kuhn and
Ludwig (1967, p. 248), and P. eurygaster has
been briefly mentioned as another louse ectopar-
asite of this monkey by Tanaka and Takefushi
(1993, p. 192); unidentified immature lice have
been reported as ectoparasites of M. fuscata at
Jigokudani Monkey Park, Shiga Heights, Nagano
Prefecture (Tanaka & Takefushi, 1993, p. 192).
Removal and ingestion of louse eggs apparently
are major motivations underlying the grooming
behavior of M. fuscata (Tanaka & Takefushi,
1993, p. 192; cf. Tanaka, 1995, p. 200; 1998, p.
1234).
Tick
At Jigokudani Monkey Park, Shiga Heights,
Nagano Prefecture, during the course of 4475 ob-
served grooming sequences, a single tick {Hae-
maphysalis longicornis) was removed from the
fur of a M. fuscata infant by its mother (Tanaka
& Takefushi, 1993, p. 189). The tick was crushed
to death by the mother but not eaten.
Q Fever Bacterium
Antibodies to Coxiella burnetii, the bacterium
that causes Q fever, were detected in sera collect-
ed from 15 of 54 M. fuscata specimens (28%) that
were live-trapped in seven prefectures in northern
and central Honshu (Ejercito et al., 1993, p. 483).
Detailed results for each prefecture are unavail-
able.
Viruses
Local variation in the incidence of antibodies
to eight viruses has been studied in three groups
of M. fuscata by Matsubayashi et al. (1992, p.
393) (Table 32). The incidence of antibodies to
two of these viruses (SA-6, Cyno-EBV) is high in
all three groups, the incidence of antibodies to
FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES
49
Table 32. Incidence {9c) of antibodies to eight vi-
ruses in three local groups of Macaca fuscata. '
Translocated groups
In-situ
Arashi-
group
yama,-
Wakasa,'
Takasaki-
Kyoto
Tottori
yama. Oita
Prefecture.
Prefecture.
Prefecture.
Honshu
Honshu
Kvushu
Virus
(n = 45)
(n = 53)
(n"= 30)
SA-6
100.0
100.0
100.0
Cyno-EBV
100.0
96.2
100.0
Measles
93.0
32.1
66.7
STLV-1
60.0
64.2
16.7
Jap. Enc.
70.5^
54.7
3.4-^
HSV-1
75.0^
1.9
0
SIVmac
4.4
0
0
SV-5
0
0
0
' Reference: Matsubayashi et al. (1992. p. 393).
- Original locality of group: virological study con-
ducted after translocation of group to outdoor enclosure
at PRIKU (Inuyama. Aichi Prefecture. Honshu) in 1981.
" Original locality of group: virological study con-
ducted after translocation of group to outdoor enclosure
at PRIKU (Inuyama, Aichi Prefecture. Honshu) in 1971.
^N = 44.
■^ N = 29.
two other viruses (SIV^,;^^. SV-5) is low in all
three groups, and the incidence of antibodies to
four viruses (measles. STLV-1. Japanese enceph-
alitis, HSV-1) is strongly variable in the three
groups. The incidence of antibodies to STLV-1
also is strongly variable (0%-ca. 9Q9c in adults)
in 41-42 local groups of M. fuscata that were
studied by Hayami et al. (1984. p. 180: cf. Hay-
ami, 1986. p. 36). The incidence of antibodies to
another virus — SV 40 — that was studied in seven
local groups of M. fuscata by Ichikawa et al.
(1985. p. 326) varied from 50% to 100% (ex-
cludes one small sample, n = 4. with 0% inci-
dence). The explanation for local variation in the
incidence of antibodies to certain viruses in M.
fuscata groups is unclear.
Antibodies to monkey B virus were absent in
8 1 M. fuscata specimens examined on Shodoshi-
ma by Tanaka et al. (1963. p. 93) and in 46 M.
fuscata specimens examined at Takasakiyama by
Endo et al. (1964. p. 174): however, they were
present in 7 of 61 captives (11.5%) originally
from Yakushima that were studied by Endo et al.
(1959, p. 229) and in 21 1 of 629 captives (33.57r)
of unknown origin that were studied by Sato et
al. (1998. p. 2(X)). Restriction fragment length
polymorphism analysis and DNA sequencing in-
dicate that the B virus that infects M. fuscata is
genetically distinct from the B viruses that infect
other macaque species (Ohsawa et al.. 2002, p.
559).
The incidence of antibodies to six viruses —
hepatitis A. HSV-1. measles. SIVm^c STLV-1,
and SV-40 — apparently tends to increase with age
in M. fuscata (Hayami et al.. 1984. p. 180: Ichi-
kawa et al.. 1985. p. 326: Hayami. 1986. p. 36:
Abe et al.. 1989. p. 88: Matsubayashi et al.. 1992,
p. 395): in two groups of monkeys tested for hep-
atitis A antibodies, antibody incidence increased
from 0% to 30% at age 1 year to 100% at age 5
years (Abe et al., 1989, p. 89). Contrastingly, the
incidence of antibodies to Cyno-EBV and SA-6
in two groups was 100% at age 1 year and re-
mained high to age ^21 year (Matsubayashi et
al.. 1992. p. 394).^
In captive groups of M. fuscata, the system of
housing apparently affects the incidence of anti-
bodies to hepatitis A (Table 33). In 43 individu-
ally housed monkeys, the antibody incidence was
2.3%. and in 48 group-housed monkeys, the in-
cidence was 70.8%.
Tanaka et al. (1963, p. 93) have reported that
M. fuscata on Shodoshima tested negative for an-
tibodies to adenoviruses (n = 26) and influenza
viruses (n = 14).
Natural History
Habitats
The two primary habitats of M. fuscata are
warm temperate evergreen broadleaf forest in the
southern part of its geographic range and cool
temperate deciduous broadleaf forest in the north-
em part of its range (Suzuki, 1965. p. 31: Uehara,
1975. p. 392: Maekawa. 1978. p. 18). The ele-
vational distribution of this species is known to
extend to 3180 m on the summit of Yarigatake
(mountain). Nagano Prefecture (S. Izumiyama,
pers. comm.. 29 Dec. 2003). During relatively
mild weather, M. fuscata has been observed in
subalpine needleleaf forest at 3050 m on Ohami-
dake (mountain), Nagano Prefecture (Izumiyama,
1987a. p. 8: 1987b. p. 42: cf. 1994. p. 478: Izu-
miyama et al.. 2003. p. 466): during snowy winter
months. M. fuscata in this part of central Honshu
moves down to the montane broadleaf forest zone
at elevations that do not exceed 1800 m (Izumi-
yama. 1987b. p. 41: cf. Ashizawa. 1983, p. 13).
A comprehensive bibliography of field studies
50
FIELDIANA: ZOOLOGY
Table 33. Relationship
fuscata.
between housing system and incidence of antibodies to hepatitis A in captive Macaca
Housing system
N
Hepatitis A
antibody
positive
Incidence
(%)
References
Individually housed
Group housed
Unspecified, possibly mixed
43
48
294
1
34
167
2.3
70.8
56.8
Chiba et al. (1980, p. 165)
Chibaetal. (1980, p. 165)
Abeet al. (1989, p. 88)
of M. fuscata (1975-1992), most of which include
habitat information, has been published by Wa-
tanabe (1993, p. 33).
Terrestriality/Arboreality
Detailed information concerning the amount of
time that M. fuscata spends on the ground and in
trees is available for only one population, which
was observed June 1995-January 1997 at Tsubaki
Wild Monkey Park, Shirahama-cho, Wakayama
Prefecture (Chatani, 2003, p. 19). In this popula-
tion, the average proportion of time that monkeys
were observed on the ground during daytime
hours was 68.3% for males and 39.8% for fe-
males. Local circumstances presumably influence
the degree of terrestriality or arboreality in M. fus-
cata. Based on research at Tsubaki Park, Chatani
(2003, p. 21) has characterized M. fuscata as
semiterrestrial — behaviorally intermediate be-
tween arboreal M. fascicularis and terrestrial M.
nemestrina.
Macaca fuscata usually sleeps in trees (Wada,
1975, p. 12; Wada & Tokida, 1985, p. 3). How-
ever, sleeping on the ground has been reported on
two small islands — Kinkazan (Takahashi, 1997, p.
60) and Kojima (Mori, 1979a, p. 43; pers.
comm.); no predators of monkeys inhabit either
of these islands.
This species reportedly swims well. In 1970, a
young male monkey swam more than 600 m from
Kojima to Torishima, an island that lies beyond
Kojima (Mito, 1980, p. 29).
Group Size and Composition
Mean group size in 117 groups of wild Japa-
nese macaques is 40.8 ± 28.95 individuals (ex-
treme values, 10-161 individuals) (Table 34; cf.
Izumiyama et al., 2003, p. 471). Groups with as
few as three or four members have been reported.
but such small groups probably are ephemeral: on
Yakushima, a group comprised of three individ-
uals— adult male, adult female, 4-year-old fe-
male— became extinct after less than 1 year of
existence (Takahata, 1991, p. 101; Maruhashi,
1992, p. 49), and at Ryozenyama, Shiga Prefec-
ture, a group comprised of four individuals, which
was observed shortly after fission of a larger
group, was not confirmed to be stable (Sugiyama
& Ohsawa, 1982a, p. 32).
Provisioned groups tend to become larger than
wild groups because favorable nutritional condi-
tions result in high birthrates and low infant mor-
tality rates. The maximum group size reported in
a provisioned group — at Takasakiyama, Oita Pre-
fecture— is 1255 individuals (Sugiyama & Ohsa-
wa, 1988).
Groups in M. fuscata are multimale and mul-
tifemale (Itani, 1954, p. 273; Takasaki & Masui,
1984, p. 310). Mean composition in 35 natural
groups is ca. 18% adult males, 32% adult females,
35% juveniles, and 15% infants (Table 35). The
mean ratio of number of infants to number of
adult females is near 0.50, which suggests that
biennial parturition is usual (see below. Repro-
duction: Birthrate).
Group size may change as a result of fission,
takeover, or fusion/extinction (Maruhashi, 1992,
p. 47). Groups that become very large often split
into two or three groups (Koyama, 1970, p. 366;
Maruhashi, 1992, p. 49); such fission reportedly
occurs between matrilines (Koyama, 1970, p.
377). Conversely, on Yakushima small groups oc-
casionally have been observed to fuse with neigh-
boring larger groups (Takahata et al., 1994, p.
320).
Mass mortality in a cluster of five neighboring
troops (65 of 1 17 troop members), including com-
plete extinction of two of the five troops, was ob-
served in western Yakushima in 1998-1999 (Han-
ya et al., 2004, p. 182).
FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES
51
Table 34. Geographic variation in group size in wild Macaca fuscata populations.
Locality (latitude)
Group size
N
Prefecture
Mean
Minimum
Maximum
References'
Aomori
Shimokita. NW (4 1=32 'N)
33
31
34
3
/
Shimokita. SW (4r09'N)
29.5
13
46
2
2, 3
Shimokita
24.5
11
42
6
4
Miyagi
Narusegawa (38°30'N)
41
—
—
1
5
Kinkazan (38'18'N)
29.3
19
55
6
5
Kinkazan (38=1 8'N)
48.7
31
79
6
4
Futakuchi (38=1 6'N)
55
—
—
1
5
Tarogawa (38°14'N)
26
—
—
1
5
Nagano
Shiga Heights (36°42'N)
25.3
15
37
6
4, 6, 7
Toyama
Kurobe (36°39'N)
23.8
10
60
18
8
Ishikawa
Hakusan (36"16'N)
41
16
86
17
4, 9, 10
Fukui
Otoumi (35°32'N)
75
—
—
1
11
Shiga
Makino-cho (35°30'N)
37.5
26
49
2
4
Ryozenyama (35°2rN)
44
—
—
1
12, 13
Hino-cho (35=01 'N)
108.3
83
134
3
4
Kanagawa
Hakone-machi (35°11'N)
32
18
46
2
4
Okayama
Katsuyama-cho (35°05'N)
112
—
—
1
4
Gagyusan (34°49'N)
120
—
—
1
4
Chiba
Takagoyama (35=10-15'N)
82.5
56
109
2
4, 14
Aichi
Shinshiro-city (34°54'N)
41.8
28
47
6
15
Osaka
Minoo-city (34°5rN)
51.5
17
86
2
4
Hiroshima
Taishakukyo (34°52'N)
29
—
—
1
4
Kagawa
Shodoshima(34°3rN)
114
52
161
3
4
Mie
Nansei-cho (34=20'N)
42
—
—
1
16
Fukuoka
Kawaradake (33=28'N)
99.5
97
102
1
4, 17
Miyazaki
Kojima(3r27'N)
22
—
—
1
18
Toi-misaki (31°22'N)
76
—
—
1
19
Kagoshima
Yakushima (30=20'N)
28.5
11
57
24
4, 20. 21
' Key to references: 1. Azuma and Ashizawa (1980). 2. Izawa and Nishida (1963). 3. Ashizawa (1979). 4. Yamagiwa
and Hill (1998). 5. Izawa et al. (2003). 6. Suzuki et al. (1975). 7. Wada and Ichiki (1980). 8. Akaza (1988). 9.
Hayashi (1970). 10. Kawai et al. (1970). 11. Watanabe (1978). 12. Sugiyama and Ohsawa (1974). 13. Sugiyama and
Ohsawa (1982b). 14. Iwano et al. (1971). 75. Kawai (1994). 16. Takasaki and Masui (1984). 17. Ikeda et al. (1973).
18. Itani and Tokuda (1958). 19. Azuma (1972). 20. Maruhashi (1980). 21. Maruhashi (1982).
Home Range Area and Group Density
Based on a survey of 32 widely distributed M.
fuscata groups. Takasaki (1981. p. 278) has
shown that home range area in this species is
strongly influenced by habitat forest type and is
positively correlated with group size (cf. Izumi-
yama et al., 2003. p. 471). In evergreen broadleaf
forest, home range area is relatively small — vary-
ing from 1.4 to 6.4 ha per group member; in de-
ciduous broadleaf forest, home range area is much
larger — varying from 9 to 79 ha per group mem-
ber.
Along the margins of home range areas, inter-
group contact behavior apparently is variable. Ag-
gression between neighboring groups reportedly
is frequent on Yakushima, less frequent on Kink-
azan. and rare or absent at Hakusan (Izawa, 1982,
p. 270; Saito et al., 1998, p. 309).
Group density on western Yakushima has been
estimated as 0.70 ± 0.43 groups/km- in undis-
turbed evergreen broadleaf forest and 1.48 ± 0.61
groups/km- in disturbed evergreen broadleaf for-
est (Hanya et al.. 2003b. p. 50).
Diet
Forest fruits, seeds, nuts, leaves, flowers,
shoots, buds, and bark generally constitute the pri-
mary natural foods of M. fuscata (Murata & Ha-
zama. 1968. pp. 5-48; Uehara, 1975, p. 395;
1977, p. 212; Maruhashi, 1980, p. 155; Iwamoto,
1982, p. 167; Koganezawa, 1983, p. 85; Watanuki
& Nakayama. 1993. p. 421; Agetsuma & Naka-
gawa. 1998. p. 279; Hanya, 2003, p. 336; Hanya
et al.. 2003a. p. 54). Other known, relatively mi-
nor dietary components include roots, grasses,
herbs, fungi, insects, crabs, spiders, mollusks,
fish, and bird's eggs (Suzuki, 1965, p. 46; Watan-
52
FIELDIANA: ZOOLOGY
Table 35. Group composition and ratios in wild Macaca fuscata (Takasaki & Masui, 1984).
Group composition
Ratine
Group
Adult
males
Adult
females
Juveniles
Infants
Totals
Locality
AdM/AdF
Inf/AdF
Shimokita
M
5
13
13
5
36
0.385
0.385
Z
7
18
10
7
42
0.389
0.389
O
4
4
3
2
13
1.000
0.500
A2
6
16
21
3
46
0.375
0.188
Shiga
B2
8
6
4
1
19
1.333
0.167
YBl
6
19
15
5
45
0.316
0.263
Hakusan
KA
9
16
14
6
45
0.563
0.375
TA
20
22
14
8
64
0.909
0.364
TB
10
20
12
4
46
0.500
0.200
O
12
18
16
7
53
0.667
0.389
Otoumi
A2
6
23
35
11
75
0.261
0.478
Takagoyama
T-Ib
6
15
28
7
56
0.400
0.467
Yugawara-machi
T
13
19
12
2
46
0.684
0.105
P
3
10
3
2
18
0.300
0.200
Minoo-city
A
6
22
41
17
86
0.273
0.773
B
1
5
8
3
17
0.200
0.600
Gagyusan
—
10
35
50
25
120
0.286
0.714
Shodoshima
I
48
37
35
9
129
1.297
0.243
KA
17
41
72
31
161
0.415
0.756
O
13
17
13
9
52
0.765
0.529
T
11
15
17
9
52
0.733
0.600
Gokashowan
—
4
13
16
9
42
0.308
0.692
Kawaradake
—
7
33
41
21
102
0.212
0.636
Kojima
—
4
6
9
3
22
0.667
0.500
Yakushima
Kojiba'
12
18
11
6
47
0.667
0.333
Nagata-todai^
7
7
9
4
27
1.000
0.571
Kannonzaki
4
4
3
2
13
1.000
0.500
Hanyama
12
12
15
6
45
1.000
0.500
Nina
7
10
5
3
25
0.700
0.300
Kawara
4
6
5
2
17
0.667
0.333
Shikamizawa
8
8
9
3
28
1.000
0.375
M
5
7
6
4
22
0.714
0.571
A
7
8
11
5
31
0.875
0.625
H
3
3
3
2
11
1.000
0.667
M
7
8
8
4
27
0.875
0.500
Totals
35
312
534
587
247
1680
—
—
Means
—
8.9
15.3
16.8
7.1
48
0.650
0.451
Group proportions
—
0.186
0.318
0.349
0.147
—
—
—
' Ko troop.
' = Todai.
abe, 1989, p. 126; Aimi & Takahata, 1994, p. 148;
Hanya, 2004, p. 61). Monkeys on Yakushima
have been observed eating frogs and lizards (Su-
zuki et al., 1990, p. 422), and on Kojima, an 11-
year-old solitary male was observed to eat a dead
shrike (Lanius bucephalus) (Kimura, 1974, p. 25).
In an upland mixed coniferous-broadleaf forest
(1000-1200 m) on Yakushima, mature leaves of
broadleaf trees were a major dietary component,
accounting for 38% of observed feeding time
(Hanya, 2004a, p. 61); consumption of leaves at
this locality apparendy increased substantially fol-
lowing a season of unusually poor fruit produc-
tion (Hanya et al., 2004, p. 184). Geophagy has
been observed on Yakushima (Maruhashi, 1980,
p. 148), on Kojima (Iwamoto, 1982, p. 157), and
at Arashiyama (Inoue, 1987, p. 104); the func-
tional significance of geophagy is unclear, but it
may serve as a buffer against gastric disorders
(Inoue, 1987, p. 104; Wakibara et al., 2001, p.
515). Feeding occupied 38% of daytime hours in
a troop studies in western Yakushima (Hanya,
2004b, p. 171).
A 5-year-old M. fuscata female at Kanbataki
Natural Park, Okayama Prefecture, reportedly ate
her dead infant a few days after the infant had
FOODEN AND AlMl: SYSTEMATIC REVIEW OF JAPANESE MACAQUES
53
died (Itoigawa, 1964, p. 13). On Yakushima, sika
deer {Cervus nippon) frequently congregate under
trees where monkeys are foraging in order to feed
on leaves, seeds, or fruit that are dropped or dis-
lodged by the monkeys (Majolo & Ventura, 2004,
p. 35).
Predators
Predation on M. fuscata by mountain hawk-ea-
gles, Spizaetus nipalensis, has been reported in
Hiroshima Prefecture and Nara Prefecture (Ha-
mada & lida, 1999, p. 18; lida, 1999, p. 125); in
the Hiroshima Prefecture attack, which occurred
on 13 July 1997, the victim was a female monkey
estimated to be 7 or 8 years old and to weigh 9.4
kg. Feral dogs formerly were abundant in Japan
and frequently preyed on monkeys (Miyamoto,
1976, p. 7), but feral dogs are now rare because
they have been severely controlled by government
action since the 1970s (Mizuno, 1995, p. 54).
Raccoon dogs {Nyctereutes procyonoides) may
have preyed on M. fuscata infants on Kojima in
1964 (Iwamoto, 1974, p. 257). The Japanese wolf,
Canis lupus hodophilax, which became extinct in
the early 1900s, may also have been a former
predator on M. fuscata.
Since 1998, more than 10,000 Japanese mon-
keys per year have been exterminated as agricul-
tural pests (Oi, 2003, p. 31).
Intergroup Dispersal
Like other macaques, M. fuscata lives in ma-
trilineal, female bonded groups (Itani, 1985, p.
602). Females remain in their natal groups, except
in cases of group fission (cf. Fukuda, 2004, p. 56).
Males over 5 years of age emigrate from their
natal groups and immigrate into neighboring or
distant groups (Norikoshi & Koyama, 1975, p. 48;
Sprague et al., 1998, p. 353); male migrations of
up to 60-100 km have been reported (Fukuda,
1982, p. 494; 1988, p. 479; Yoshimi & Takasaki,
2003, p. 71). Because spermatogenesis in M. fus-
cata begins ca. age 4-6 years (Nigi, 1975a, p. 51;
Nigi et al., 1980, p. 237; see below. Reproduc-
tion), close inbreeding is avoided.
Group tenure of males varies from 0.2 to 5.3
years (mean = 2.9 years) on Kinkazan, from 0.2
to 5.7 years at Arashiyama (Norikoshi & Koyama,
1975, p. 55), and from 1.0 to 9.5 years (mean =
2.8 years) on Yakushima (Sprague et al., 1998, p.
352).
Reproduction
Seasonality
Reproduction is strongly seasonal in M. fusca-
ta, with matings restricted to autumn and winter
and births restricted to spring and summer (Kawai
et al., 1967, p. 40; Nozaki et al., 1992, p. 301).
Within the spring-summer period, the timing of
births apparently varies geographically (Fooden &
Aimi, 2003, p. Ill), and this presumably also ap-
plies to the timing of matings within the autumn-
winter period. In in-situ troops that inhabit the lat-
itudinal zone between ca. 31°20'N and 38°20'N,
the mean birth date tends to become earlier as
latitude increases (Figs. 21, 22); this pattern of
variation may function to provide more time for
infants born at higher latitudes to achieve an ad-
equate level of development before the onset of
subsequent poor winter feeding conditions, which
generally begin earlier at higher latitudes than at
lower latitudes.
The above relationship between mean birth date
and latitude apparently does not apply to troops
that inhabit localities at the geographic and eco-
logical extremes of the distribution of M. fusca-
ta— namely, Yakushima, ca. 30°20'N, and Shi-
mokita Peninsula, ca. 4riO'N (Fig. 22); on Yak-
ushima, the mean birth date is earlier than ex-
pected, and on Shimokita Peninsula, it is later
than expected. The mild winters on Yakushima
may be less of a factor in infant survival than
elsewhere in Japan. Conversely, winters on Shi-
mokita Peninsula last so long that earlier births
might result in infants being born before anteced-
ent poor winter feeding conditions have ended.
Troops that have been translocated to new lat-
itudes in the northern hemisphere generally retain
approximately the same mean birth date that they
manifested at their original localities (Fooden &
Aimi, 2003, p. 111). Contrastingly, in one group
that was translocated to the southern hemisphere
(Tasmania), the mean birth date shifted approxi-
mately 6 months forward; this presumably was in
response to the reversed seasons. Laboratory cap-
tives kept under conditions of constant artificial
day length and temperature retained their original
ovulatory and birth cycles for at least 12 years
(Nozaki, 1991, p. 104; Nozaki et al., 1992, p.
54
FIELDIANA: ZOOLOGY
In-situ troops
Translocated troops
130°
135'
140*'
Fig. 21. Distribution of Macaca fuscata troops in Japan for which birth seasonality data are available (for details,
see Fooden & Aimi, 2003, p. 1 15).
In-situ troops. Localities with complete birth seasonality data: 1. Yakushima: Nina-A. 2. Toimisaki; 3. Kojima; 4.
Takasakiyama; 5. Nametoko; 7. Kochi; 8. Taishakukyo; 9. Gagyusan; 10. Kanbanotaki; 13. Funakoshiyama; 14.
Rosando; 15. Choshikei; 17. Minoo-city-A; 18. Arashiyama; 23. Hagachi; 25. Takagoyama-S; 26. Takagoyama-A;
29. Jigokudani; 30. Kinkazan A; 31. Shimokita Peninsula: Wakinosawa. Localities with incomplete birth seasonality
data (not plotted in Fig. 21): 11. Katsuyama-cho; 16. Kaminada; 19. Otoumi; 20. Ryozen-A; 24. Hakone-machi.
Translocated troops. 6. Miyajima-cho; 12. Takeno-cho; 21. Ohirayama; 22. Okinoshima; 27. Takaosan; 28. Hou-
tosan.
FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES
55
1 1 1 1 1 1 1 1
■po das 6nv inp unp Ab|^ Jdy JBI/^ qej
Inside Toimisaki- Outside Toimisaki-
Kinkazan Kinkazan
latitudinal zone latitudinal zone
Earliest birth date • o
Latest birth date ■ d
Mean birth date ♦ O
■ A . A
■ * / •
_ ■ ^ 1 m m.
^ 1 .
■ I » ^^^ ^n^ . 1
■ iV •
▼ / •
_ ■ — • / * ^ — _ m
■ ■
^ 1 .
7^
■ A. / A
CM
-"it
O
CO
CO
CO
co ^^
Z
M
0>
CD
CO 2.
a>
XJ
3
(S
in
CO
CO
CO
CO
CM
CO
O
O
CO
o
00
CNI
O
CD
CNI
O
CM
O
CNJ
CM
O
O
CM
O
CO
O
CO
O
CO
O
O
CNI
O
O
O O
OO CD
O
•ou Abq
56
FIELDIANA: ZOOLOGY
302). Experimental manipulation of photoperiod
had little or no effect on ovulatory cycles of cap-
tives, but simultaneous manipulation of photope-
riod and temperature resulted in reversed cycles
in two of four experimental monkeys.
Sexual Maturation
Although M. fuscata males as young as age 1 .5
years have been observed to mount females (Tak-
ahata, 1980, p. 325; Eaton et al., 1986, p. 144;
Mehlman & Chapais, 1988, p. 202), ejaculation
in this species normally does not occur until age
4.5 years (Hanby et al., 1971, p. 125; Wolfe,
1978, p. 62; Takahata, 1980, p. 325; Inoue et al.,
1991, p. 203); one exceptional case of ejaculation
at age ca. 3.5 years has been reported by Chapais
and Mignault (1991, p. 173). Descent of the testes
into the scrotum in various troops of free-ranging
M. fuscata has been reported to occur at age ca.
4.5 years (Hiraiwa, 1981, p. 315), at age ca. 5.5
years (Nigi et al., 1989, p. 572), or even after age
6 years (Mori, 1979b, p. 381); Nigi et al. (1989,
p. 574) have suggested that sexual maturation in
males may be accelerated by artificial feeding.
Histologically, sperm are rarely detected in the
testes of monkeys at age ca. 3.5 years (Inoue &
Hayama, 1963, p. 76; Nigi, 1975a, p. 51; Matsu-
bayashi & Mochizuki, 1982, p. 895), but they are
commonly seen in males at age 4.5 years (Nigi,
1975a, p. 51; Nigi et al., 1980, p. 233; Matsubay-
ashi & Mochizuki, 1982, p. 897). By age 5 years,
almost all males have emigrated from their natal
troops (Sugawara, 1975, p. 354; Hasegawa & Hir-
aiwa, 1980, p. 136; Fukuda, 1982, p. 493; Sugi-
yama & Ohsawa, 1982b, p. 255; Matsumura,
1993; Sprague et al., 1998, p. 353; Yoshimi &
Takasaki, 2003, p. 72) (see above. Natural His-
tory: Intergroup Dispersal). Full sociosexual ma-
turity does not occur until age >8.5 years, when
the frequency of mating activity is approximately
five times as great as at younger ages (Takahata,
1980, p. 325); by this age, males have developed
mature or near-mature sexual skin coloration (Ma-
sui et al., 1973, p. 237; Mori, 1979b, p. 381; Hir-
aiwa, 1981, p. 315; Maruhashi, 1982, p. 320).
Menarche (first menstruation) in M. fuscata fe-
males usually occurs at age ca. 3.5 years (Nigi,
1975b, p. 208; Mori, 1979b, p. 381; Hasegawa «&
Hiraiwa, 1980, p. 136; Hamada et al., 1999, pp.
441, 449; cf. Hazama, 1964, p. 89). At about the
same age, characteristic para-anal pubertal swell-
ings become evident (Wolfe, 1979a, p. 412; Tak-
ahata, 1980, p. 304; Maruhashi, 1982, p. 320; Oi,
1988, p. 13; cf. Pocock, 1906, p. 558; Mori et al.,
1997, p. 562); these are succeeded by subcaudal
or paravulval pubertal swellings during the mating
seasons of the next 2-3 years. Females typically
begin to display estrous behavior at age ca. 3.5
years or ca. 4.5 years (Wolfe, 1979a, p. 412; Hir-
aiwa, 1981, p. 311; Chapais & Mignault, 1991, p.
173; Pavelka, 1993, p. 70). In provisioned troops,
females generally give birth to their first infants
at age ca. 5 years or age ca. 6 years (cf. Fedigan,
1991b, p. 145; Itoigawa et al., 1992, p. 57; Koya-
ma et al., 1992, p. 37); known extreme ages of
primiparous females are ca. 4 years and ca. 16
years (Watanabe et al., 1992, p. 8). Provisioning
of troops apparently tends to reduce the mean age
of primiparity of troop members (Sugiyama &
Ohsawa, 1982a, p. 37; Fukuda, 1988, p. 481;
Watanabe et al., 1992, p. 8).
Sexual Skin
During the mating season, the bare skin on the
face, nipples, and perineum of M. fuscata be-
comes intensely reddish in adults and, to a lesser
degree, also in adolescents. In females, this red-
dening varies cyclically during the mating season,
increasing during estrus intervals and decreasing
during interestrus intervals (Tokuda, 1963, p. 14;
Nigi, 1975b, p. 21 1; Wolfe, 1984c, p. 146; Taka-
hata et al., 1995, p. 170); estrus intervals are also
marked by vaginal discharge of a distinctively
odoriferous secretion (Enomoto, 1974, p. 353;
Takahata et al., 1995, p. 170). Unlike adolescent
females (see above, Sexual Maturation), adult fe-
FiG. 22. Latitudinal variation in birth seasonality in in-situ troops oi Macaca fuscata (for documentation, see Fooden
& Aimi, 2003, p. 1 15). Seasonality in 18 troops distributed within the Toimisaki-Kinkazan latitudinal zone (ca. 3r20'-
38°20'N) is compared with seasonality in troops on Yakushima (Nina-A, 30°2rN) and Shimokita Peninsula (Waki-
nosawa, 41°08'N). Regression equation: y = -10.07x + 493.4; r = -0.764, F = 16, P < O.OI. (Note: In 2003, data
for which are not included in this graph, the mean date of 16 births at Wakinosawa was 30 April, and extreme birth
dates were 10 April and 28 May [S. Matsuoka, Working Group for Monkeys in the Shimokita Peninsula, pers.
comm.]).
FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES
57
Table 36.
Menstrual
cycle length in Macaca fuscata samples (mean ~ 27.1
days, n ~ 593
cycles).
Publication
date
Mean cych
length
(days)
i Standard Extreme
deviation cycle lengths Number of
(days) (days) females
Number of
cycles
Reference'
1980
26.52
Arashiyama-B: free-ranging provisioned troop
9.1 8-53' —
73
;
1989
27.5
Ehime University
7.8 19-47 26
94
2
1975
1977
1977
Japan Monkey Centre (mean ~ 26.2 days, n ~ 315 cycles)
26.3 5.4 < 15-35^ 10 -275
29.5 1.7 28-32 4 4
25.3 2.8 19-31 — 36
3
4
5
1977
1992
Primate Research Institute, Kyoto University (mean = 30.2 days, n
28.5 3.0 23-36 7
31.7 3.2"^ — —
= 76 cycles)
36
40
6
7
1979
28.1
Tokai University
4.2 21-40 6
35
8
' Key to references: 7. Takahata (1980, p. 311). 2. Hamada et al. (1989, p. 188). 3. Nigi (1975b, p. 210). 4. Ando
et al. (1977, p. 274). 5. Nigi (1977, p. 246). 6. Oshima et al. (1977, p. 101; cf. Aso et al., 1977, p. 747). 7. Nozaki
et al. (1992, p. 304). 8. Enomoto et al. (1979, p. 564).
' Mode = 26 days.
-^ Excludes cycles >53 days; intervening bleeding possibly missed.
'• Excludes 14 cycles, duration >40 days.
^ Standard error = 0.5.
males do not exhibit perineal swellings (Nigi,
1975b, p. 211; Wolfe, 1984c, p. 146), although
they do undergo cyclical reddening of the peri-
neum. In ovariectomized females, reddening of
the sexual skin can be induced by injection of
estrogen (Nigi & Ando, 1975, p. 266).
In adult males, the intensely red coloration of
the sexual skin — including the face, perianal area,
and scrotum — persists more or less continuously
throughout the mating season (Alexander, 1970,
p. 279; Enomoto, 1974, p. 353; Wolfe, 1979a, p.
413; Maruhashi, 1982, p. 319). Concurrently dur-
ing the mating season, adult males carry their
short tails erect, which increases the visibility of
the bright red perineal sexual skin (Wolfe, 1981a,
p. 26). Near the beginning of the mating season,
increased redness of the sexual skin and seasonal
erection of the tail are known to be associated
with increased testosterone levels (Rostal et al.,
1986, p. 459).
Menstrual Cycle
During the mating season, menstrual cycle
length averages ca. 27.1 days (Table 36), and the
duration of menstrual flow averages 3-5 days
(Nigi, 1975b, p. 210; Aso et al., 1977, p. 747;
Oshima et al., 1977, p. 101; Hamada et al., 1989,
p. 188). A female's peak estradiol value, which is
followed approximately 1 day later by ovulation,
occurs near the middle of the menstrual cycle
(Nomura et al., 1972, p. 475; Ando et al., 1976,
p. 92; Ando et al., 1977, p. 274; Nigi & Torii,
1983, p. 413) During the nonmating season, men-
strual cycle length becomes irregular, and men-
struation may be completely suppressed (Tokuda,
1963, p. 11; Nomura et al., 1972, p. 474; Nigi,
1975b, p. 210; Hamada et al.. 1989, p. 188).
Estrus
Available information indicates that, during the
mating season, a mature M. fuscata female will
exhibit 0-6 (mean, ca. 2) discrete periods of sex-
ual proceptivity and receptivity — i.e., estrus (Ta-
ble 37). The known duration of these estrous pe-
riods, which are not synchronized among female
troop members, varies from 1 to 92 days (mean,
ca. 10 days); the intervals between estrous periods
also are highly variable (Wolfe, 1984c, p. 150;
Pavelka, 1993, p. 96). In a recent study of the
relationship between hormonal cycles and sexual
behavior in a seminatural population of M. fus-
cata, female attractivity and proceptivity were
58
FIELDIANA: ZOOLOGY
Table 37. Frequency and duration of estrous periods in mature Macaca fuscata females.
Number of estrous periods
Estrous period duration (days)
Observation .
per female
per matmg
season
Refer-
Locality
period
Mean ± SD
Extremes
N
Mean ± SD
Extremes
N
ences'
Arashiyama
1975-77
2.6 ± 1.5
0-6
87
13.6 ± 12.2
1-92
87
1
Arashiyama
1977-78
2.8 ± 1.5
—
71
-8.5
—
198
2
Arashiyama
1979-85
1.9 ± 0.9
1-4
156
8.9 ± 8.4
1-42
156
3
Kojima
1950s
—
—
—
9.3 ± 3.1
6-13
26
4
Shiga A
1967-73
-1.7
1-5
-77
-10.7
1—75
-264
5
South Texas-
1973-74
2.5 ± 1.1
—
—
8.3 ± 9.8
—
—
6
' Key to references: 7. Takahata (1980, pp. 311, 314). 2. Wolfe (1984c, p. 151). 3. Huffman (1991b, p. 104). 4.
Tokuda (1961-62. p. 13). 5. Tokita and Hara (1975, p. 30). 6. Wolfe (1979b, p. 527).
- Translocated from Arashiyama.
found to increase during the follicular and peri-
ovulatory phases of the hormonal cycles (O'Neill
et al., 2004b, p. 359).
At Arashiyama, estrus in females with infants
(born ca. 6 months before the mating season) was
delayed by approximately 1 month relative to es-
trus in females without infants (Takahata, 1980,
p. 317), and on Kinkazan, estrus was rare in fe-
males with infants (one estrous season in 19 fe-
male-years) (Takahashi, 2002, p. 145). In the Shi-
ga A troop, most females reportedly exhibited
clear estrus only in alternate years (Enomoto,
1975. p. 275; 1978, p. 284).
On Kinkazan, fruit abundance during the mat-
ing season strongly affected the observed estrus
rate (Takahashi, 2002, p. 148). At Arashiyama,
dominance rank influenced estrus duration; the to-
tal annual duration of estrus in high-ranking fe-
males (mean = 58.8 days, n = 22) was approxi-
mately twice as great as that in lower-ranking fe-
males (mean = 28.2 days, n = 32) (Takahata,
1980, p. 324).
Most conceptions in M. fuscata occur during a
female's first estrous period of the mating season
(Takahata, 1980, p. 311; Huffman, 1993, p. 80;
Maruhashi & Takasaki, 1996, p. 154). However,
even after conception, females usually exhibit one
or more additional estrous periods — with postcon-
ception copulations — during the remainder of the
mating season (Hanby et al., 1971, p. 137; Taka-
hata, 1980, p. 311; 1999, p. 148; Wolfe, 1979b,
p. 527; 1984c, p. 151; Huffman, 1993, p. 80).
Male sexual activity in M. fuscata is more or
less continual during the mating season and does
not exhibit estrus-like periodicity. Two cases of
unseasonably late estrous periods followed by un-
seasonably late births suggest that the mating ac-
tivity of males may be induced by the presence
of estrous females (Kawai, 1966, p. 392).
Consortship
During the mating season, males and estrous
females in natural and seminatural groups form
consortships — temporary associations of varying
duration in the course of which copulations usu-
ally occur (Imanishi, 1963, p. 77; Itani, 1963, p.
51; Eaton, 1976, p. 103; D'Amato et al., 1982, p.
228; Gouzoules & Goy, 1983, p. 41; Yamagiwa,
1985, p. 114; Huffman, 1991c, p. 200; cf. Perloe,
1992, p. 296). Consortships may be initiated by
either sex (Itani, 1963, p. 51; Tokuda, 1963, p.
16; Stephenson, 1973, pp. 67-71; 1975, p. 74;
Wolfe, 1979b, p. 527; 1984c, p. 148; Xiong &
Wang, 1991, p. 14; Soltis et al., 1997b, p. 733;
1997a, p. 744; 1999, p. 272); however, females
and males that form consortship pairs usually are
not individuals that are closely affiliated during
the nonmating season (Enomoto, 1975, p. 278;
Fedigan & Gouzoules, 1978, p. 493; Baxter &
Fedigan, 1979, p. 451; Takahata, 1982b, p. 100).
In the Arashiyama B troop, the mean (±SD) du-
ration of 168 uninterrupted consortships was 1.6
±1.1 days (extremes, 1-7 days) (Huffman, 1993,
p. 80).
Troop females often form consortships with
nontroop solitary males, which tend to approach
troops during the mating season (Nishida, 1966,
p. 171; Kawai et al., 1968, p. 10; Kawanaka,
1973, p. 149; numerous subsequent authors).
Dominant males frequently attempt to disrupt the
consortships of subordinate males (Tokuda, 1963,
p. 29; Furuichi, 1985, p. 230; Huffman, 1987, pp.
FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES
59
Table 38. Reported results concerning relationship between male dominance rank and reproductive success in
natural and seminatural groups of Macaca fitscata.
Reports that variables are
positively correlated
Reports that variables are not
positively correlated
-male dominance rank and copulation success
Enomoto (1974. p. 367)
Stephenson (1975. p. 92)
Takahata (1982b. p. 105)
Xiong and Wang (1991. p
16)
Natural groups: Variables
Nishida(1966. p. 169)
Stephenson (1975. p. 85)
Huffman (1987. p. 231)
Soltis et al. (2001. p. 488)
Matsubara (2003. p. 1062)
Natural groups: Variables — male dominance rank and fertilization success
Soltis et al. (2001. p. 488) Huffman (1987. p. 231 )
Seminatural groups: Variables — male dominance rank and copulation success
Eaton (1971, p. 9) Eaton (1974. p. 292)
Hanbv et al. (1971. p. 132) Soltis et al. (1997a. p. 741)
Stephenson (1975. p. 98)
Inoue et al. (1991. p. 204)
Seminatural groups: Variables — male dominance rank and fertilization success
Inoue et al. (1990. p. 567) Inoue et al. (1991. p. 204)
Soltis et al. (1997a. p. 742)
223-224: 1991b. p. 106: 1992, p. 83): presumably
as a consequence, consortships involving domi-
nant males usually occur near the center of a
troop, and those involving lower-ranking males,
or nontroop males, usually occur at or beyond the
periphery of a troop (Imanishi. 1963, p. 77: Fu-
ruichi. 1985. p. 230: Sprague. 1994. p. 185). Each
sexually mature male or female is likely to have
more than one consortship partner during the
course of a single mating season (Tokuda. 1963.
p. 36: Eaton, 1976. p. 103: Wolfe. 1986. p. 271:
Inoue et al.. 1993. p. 507: Soltis et al.. 1999. p.
270: 2001. p. 489: Matsubara & Sprague. 2004.
p. 904): in a seminatural group that included 79
sexually mature females, the mean (±SD) number
of male partners per female during one mating
season was 4.78 ± 3.68 (extremes. 1-15) (Gou-
zoules & Goy. 1983. p. 41). Polyandry may ben-
efit females by reducing infant harassment and in-
fanticide (Soltis et al.. 2000. p. 197: Soltis. 2002.
p. 192: see below. Infant Mortality).
Copulatory Behavior
M. fuscata generally is a multimount ejacula-
tor — i.e.. each ejaculatory copulation usually con-
sists of a series of mounts separated by brief (ca.
30-second) dismounts (Tokuda. 1963, p. 28: Han-
by et al.. 1971, p. 127: Hanby & Brown. 1974, p.
162: Enomoto, 1974, p. 363: Stephenson, 1975.
p. 74: Wolfe. 1978, p. 60: Xiong & Wang. 1991.
p. 14: Pavelka. 1993. p. 101: Troisi & Carosi,
1998. p. 1263). The mean number of mounts per
copulatory sequence reported in various natural
and seminatural troops vaiies from 8.8 to 18.9
(extremes. 1-59), and the mean duration of cop-
ulatory sequences varies from 6.2 to 13.0 minutes
(extremes. l->60 minutes): the mean number of
intromissive thrusts per mount vaines from 1.7 to
3.7.
As in M. nmlatta. a M. fitscata male usually
mounts a female dorsoventrally by gripping the
female's waist and shanks with his hands and feet,
respectively (Tokuda. 1963. p. 26: Hanby et al.,
1971, p. 126: Hanby & Brown. 1974, p. 156:
Wolfe, 1978, p. 58). Copulation occurs both in
trees and on the ground (Yotsumoto. 1976. p.
202). During the daytime, copulation is most fre-
quently observed early in the morning (Hanby et
al.. 1971, pp. 127-131: Oda & Masataka, 1995,
p. 133: Vasey, 1998b, p. 583). but optically as-
sisted night viewing and fresh copulatory plugs
observed at dawn indicate that copulations also
occur at night (Inoue et al., 1991, p. 206: Soltis
et al.. 1997a, p. 744).
Dominance Rank and Reproductive Success
Equivocal results have been reported concerning
the relationship between male dominance rank and
reproductive success in M. fitscata (Table 38). This
may indicate variation in this relationship among
60
FIELDIANA: ZOOLOGY
the groups studied and/or variation in the meth-
odology of observers (cf. Takahashi, 2004, p. 101).
In females, dominance rank and birthrate were
found to be positively correlated in two studies (see
below. Birthrate) and were found to be uncorrelat-
ed in four studies. In a small captive group of M.
fuscata, the dominant female successfully restricted
the copulation and fertilization of a subordinate fe-
male (Rendall & Taylor, 1991, pp. 323, 325).
Inbreeding
In natural populations of M. fuscata, the op-
portunity for close inbreeding is minimized by the
strong tendency for young males to leave their
natal troops before they reach full sociosexual
maturity (see above. Sexual Maturation). Some
adolescent males that have not yet emigrated may
exhibit positive sexual behavior toward closely re-
lated females, but these advances generally are
rebuffed by the females (Enomoto, 1974, p. 369).
Aunt-nephew matings are the most closely inbred
pairings with ejaculatory copulations that have
been reported in natural troops (Koyama, 1970, p.
357; Enomoto, 1978, p. 290; cf. Imanishi, 1961,
p. 119; Takahata et al., 2002, p. 406).
Although male emigration is impossible in con-
fined seminatural populations, close matrilineal
inbreeding also is rare in such groups (Eaton,
1971, p. 9; Hanby & Brown, 1974, p. 163; Wolfe,
1977, p. 168; Baxter & Fedigan, 1979, p. 454;
Pavelka, 1993, p. 109; Soltis et al., 1997a, p. 739);
father-daughter matings and matings between pa-
ternal half siblings have been observed in semi-
natural groups (Inoue et al., 1990, p. 568; 1992,
p. 134). As in natural troops, rejection by females
probably is a major factor in the inhibition of ma-
trilineal inbreeding in seminatural groups (Soltis
et al., 1999, p. 270). Homosexual behavior ap-
parently also is rare between close matrilineal rel-
atives, at least in seminatural groups (Fedigan &
Gouzoules, 1978, p. 493; Mehlman & Chapais,
1988, p. 202; also see below, Nonreproductive
Sexual Behavior).
No serial mounts were observed between a M.
fuscata mother and her adult son that were ex-
perimentally caged together during three mating
seasons (Itoigawa et al., 1981, p. 502).
Hybridization
Breeding between M. fuscata and other ma-
caque species has occurred both in captivity and
under free-ranging conditions. In zoos, M. fuscata
reportedly has hybridized with M. cyclopis and M.
silenus (International Zoo Yearbook, 1963, p. 224;
1 97 1, p. 265). Under free-ranging conditions, hy-
bridization with an introduced population of M.
mulatta has been observed, and hybridization with
an introduced population of M. cyclopis has been
inferred (see below). Judging from available evi-
dence, females and males in free-ranging M. fus-
cata troops spontaneously interbreed with other
macaque species when given the opportunity.
At Arashiyama, near Kyoto, two estrous fe-
males— members of a local free-ranging troop of
M. fuscata — climbed over an electrified fence that
enclosed a small imported group of M. mulatta,
copulated with M. mulatta males, climbed out of
the enclosure, and subsequently gave birth to hy-
brid infants (Wolfe, 1981b, p. 131). One of the
hybrid infants, a female bom in 1968, was per-
mitted to remain in the M. fuscata troop; when
she reached adulthood, her size and dorsal pelage
color apparently were intermediate between that
in M. fuscata and M. mulatta, her tail length ap-
parently was similar to that in M. mulatta (cf.
Fooden, 2000, p. 30), and her estrous perineal
sexual skin apparently was similar to that in M.
fuscata. In 1972 and 1973, this hybrid gave birth
to two daughters; as adults, the daughters (0.75
M. fuscata, 0.25 M. mulatta) resembled their
mother in size and dorsal pelage color, but their
tail length was intermediate between that of their
mother and that of M. fuscata.
In Wakayama Prefecture, south of Osaka, an
imported group of M. cyclopis escaped from a lo-
cal menagerie in 1955 and became established in
a nearby forest south of Ooike village (Kawamoto
et al., 1999, p. 53; 2001, p. 13). A recent study
of blood proteins and mtDNA in the Ooike pop-
ulation indicates that M. fuscata males from
troops in adjacent areas have freely hybridized
with females in the introduced M. cyclopis pop-
ulation; approximately 25% of the current Ooike
population gene pool is estimated to be of M. fus-
cata origin. The tail length reported in a young
male F, hybrid (age 4-5 years) is 290 mm, which
is approximately intermediate between tail length
in M. fuscata and that in M. cyclopis (see above.
External Measurements and Proportions; Fooden
& Wu, 2001, p. 13).
At the southern tip of Boso Peninsula, Chiba
Prefecture, M. fuscata males apparently have hy-
bridized with introduced individuals or groups of
M. fascicularis, M. cyclopis, and M. mulatta (Ai-
zawa & Hagiwara, 2001, p. 4).
FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES
61
Table 39. References' concerning nonreproductive sexual behavior reported in natural and seminatural groups
of Macaca fuscata.
Nonreproductive
mounts-
Masturbation
Locality
;/
^
-j;
z/z
^
'
Natural groups
Arashiyama
7.2..
^,4
5
6
3
Kojima
•)
1,5
7
Minoo-ciiy
2
8
Miyajima-cho"
5
Shiga
9
9
9
9
9
Takasakiyama
9
9
9
9
9. 10
Yakushima
11
Seminatural
groups
Arashiyama West. U.S. .A.
12, 13.
14
14. 15
12
15
Beaverton. U.S. .A.
16,17
16
16,
17,
18
16.19
Calgary Zoo. Canada
20
20
Ca\ riglia Park. Italy
21
St. Hyacinthe. Canada^
22
Universite de Montreal,
Canada
23,24
23
' Ke\ to references: /.
Takahata (1980.
pp. 306.
315. 323).
2. Takahata
(1982b. p.
104).
3. Wolfe
(1984c. pp. 150,
151). 4. Wolfe (1986. pp. 268. 271 ). 5. Stephenson (1973. pp. 63. 66. 70). 6. Takenoshita (1998. p. 365). 7. Stephenson
(1975. p. 92). 8. Perloe (1992. p. 300). 9. Eonomoto (1974. pp. 354. 364). 10. Itani (1959. p. 86). //. Thomsen
(2000. p. 213); Soltis et al. (2001. p. 487): Domingo-Roura et al. (2004. p. 34). 12. Fedigan and Gouzoules (1978,
p. 493). 13. Baxter and Fedisan (1979. p. 454): Wolfe (1979b. pp. 526. 531: 1984a, p. 23^5); Yoshida (1988. p. 58);
Pavelka (1993. p. 108). 14. Gouzoules and Gov (1983. p. 42); O'Neill et al. (2004a. p. 29). 15 Wolfe (1978, pp. 58,
63). 16. Hanby et al. (1971. pp. 126. 132. 133. 137). 17. Eaton (1978, p. 49). 18. Hanbv (1974. pp. 838. 843). 19.
.■\lexander (1970. p. 279). 20. Rendall and Tavlor (1991, p. 324). 21. Lunardini (1989. p.'l83); Corradino (1990, pp.
356. 358). 22. Vasey (1996, pp. 543. 550; 1998b. pp. 582. 585. 590): Chapais et al. (1997. p. 1096); Vasev et al.
(1998, p. 392); Vasev and Gauthier (2000. pp. 20, 22). 23. Mehlman and Chapais (988. p. 202). 24. Chapais and
Mignauh (1991. pp. 174. 175).
- Tabulated references to homosexual mounts pertain exclusively to mount series; references to single-mount events,
which usually are not sexually motivated, are excluded.
'This population was translocated from Shodoshima in 1962 (Kawai et al.. 1967. p. 47).
- This population was translocated from the Universite de Montreal in 1992 (Chapais, 1986, p. 409; Chapais et al.,
1997. p. 1091; Vasey. 1998b. p. 583).
Nonreproductive Sexual Behavior
Homosexual and autosexual behaviors have
been observed in natural and seminatural groups
of A/, fuscata (Table 39); these behaviors include
female/female mounts, female/male mounts,
male/male mounts, and masturbation by females
and males. As indicated below, published data
sets concerning these behaviors often are frag-
mentary or incongruent and therefore are some-
v^hat difficult to compare.
In natural groups (Table 39), female/female
consortships. including reciprocal mounts, have
been reported at Arashiyama (47*!^ of 94 sexually
mature females in 1976^1977: TT^ of 88 sexually
mature females in 1977-1978). Kojima (rarely
observed). Minoo. Shiga, and Takasakiyama (no
frequency data for the last three localities); at Ara-
shiyama. the homosexual estrous p>eriods of a fe-
male usually did not overlap with her heterosex-
ual estrous periods (ca. 679^ of observed estrous
periods). In addition to mounting other females,
females also have been observed to mount males
during the course of heterosexual consortships;
this behavior has been reported at Arashiyama
(lO'v^ of 87 observed heterosexual pairs), on Mi-
yajima (14% of 49 observed heterosexual pairs),
at Shiga and Takasakiyama. but never on Kojima
(>58 observed heterosexual pairs). Male/male
consortships and ejaculatory mounts, including
reciprocal mounts, have been reported at Arashi-
yama (six males in two large troops [>300 indi-
viduals]). Shiga, and Takasakiyama: at Arashiya-
ma. two of six males that participated in homo-
sexual mounts also were observed to engage in
heterosexual activity during the same mating sea-
son. Female masturbation (stimulation of clitoris
manually or with inanimate object) has been re-
ported at Arashiyama (167r of 111 females). Shi-
ga, and Takasakiyama. Male masturbation, includ-
62
FIELDIANA: ZOOLOGY
ing masturbation by troop leaders, has been re-
ported at Kojima, Shiga, and Takasakiyama and
on Yakushima.
In seminatural groups (Table 39), at Arashiya-
ma West, Calgary Zoo, Cavriglia Park, Universite
de Montreal, and St. Hyacinthe, homosexual con-
sortships and mounts were reciprocally performed
by 51-100% of sexually mature females, whereas
at Beaverton the reported mean frequency of fe-
male homosexual mounters was only 9.7%; al-
most all females that participated in homosexual
mounts also participated in heterosexual mounts.
At Arashiyama West, O'Neill et al. (2004a, p. 29)
have discovered that female homosexual
mounts — like heterosexual mounts — are restricted
to the follicular and periovulatory phases of the
ovarian cycle. Females that were engaged in ho-
mosexual consortships have been observed to re-
ject the sexual solicitations of intruding males
(Vasey, 1998b, p. 590); homosexual consortships
apparently are rare or absent between closely re-
lated females (Wolfe, 1979b, p. 531; Chapais &
Mignault, 1991. p. 175; Chapais et al., 1997, p.
1096; Vasey & Gauthier, 2000, p. 20). During the
course of heterosexual consortships, females have
been observed to mount males at Arashiyama
West (31% of 1 14 copulations in 1973-1974; 42%
of 79 sexually active females in 1977-1978), Bea-
verton (ca. 40% of 37-58 sexually active fe-
males), and Universite de Montreal (25% of eight
sexually active females). Male/male mount series
have been reported at Arashiyama West (ca. 1%
of 702 consortships during three mating seasons)
and Beaverton (ca. 15% of 22-59 sexually active
males during four mating seasons; extremes, 4-
127 male/male mount series per mating season).
Female masturbation in seminatural groups has
been reported only at Calgary Zoo (three of three
sexually mature females). Male masturbation, in-
cluding manual and oral stimulation of penis, has
been reported at Arashiyama West (2 of ca. 15
sexually mature males) and Beaverton (13 of 26
sexually active males; 27 masturbations, 12 ejac-
ulatory). In individually caged M. fuscata males
at PRIKU, the mean daily frequency of mastur-
bation during sampled mating-season months of
October and December (0.63 masturbations/day)
was nearly four times as great as during sampled
birth-season months of April and June (0.17 mas-
turbations/day) (Matsubayashi, 1974, p. 254).
Two tentative generalizations are suggested by
the available data: (1) The frequency of some or
all of the above nonreproductive sexual behaviors
probably varies locally and annually in M. fus-
cata. (2) The frequency of female homosexual be-
havior in M. fuscata probably exceeds the fre-
quency of male homosexual behavior (cf. Vasey,
1995, p. 179).
Various interpretations of nonreproductive sex-
ual behaviors have been published. Eaton (1978,
p. 48) and Wolfe (1979b, p. 532; 1984a, p. 235;
1984c, p. 152; 1986, p. 273) have noted that var-
iation in the frequency of female and/or male ho-
mosexual behavior apparently is related to varia-
tion in the sex ratio of mature troop members; this
is interpreted to indicate that homosexual behav-
ior in M. fuscata may be a response to an inade-
quate number of suitable heterosexual partners
(cf. Fedigan & Gouzoules, 1978, p. 494; Rendall
& Taylor, 1991, p. 326; Takenoshita, 1998, p. 366;
Vasey & Gauthier, 2000, p. 22). Gouzoules and
Goy (1983, p. 47) have suggested that mounting
by females may be a learned behavior that is de-
pendent on parturition or pregnancy status. Vasey
(1996, p. 550; 1998a, p. 417; 1998b, p. 595; Va-
sey et al., 1998, p. 395; Vasey & Gauthier, 2000,
p. 23) and coworkers have proposed that female/
female consortships are motivated by sexual at-
traction and gratification and that this behavior
persists because it is selectively neutral.
Wolfe (1984c, p. 152) has suggested that female
masturbation may be a learned behavior that is so-
cially transmitted within matrilines, whereas Ren-
dall and Taylor (1991, p. 326) indicated that this
behavior may be a result of restricted access to a
suitable mating partner Thomsen (2000, p. 213)
has proposed that male masturbation may function
to improve ejaculate quality by removing sperm
that have begun to deteriorate; this interpretation
would not be applicable to female masturbation.
Gestation
In a composite sample of 73 M. fuscata preg-
nancies, mean duration of the gestation period is
171.7 days (Table 40); although the standard de-
viation of this composite mean is indeterminable,
extreme durations are 157 and 188.5 days. For
infants of known gestation length, sex has been
reported for only two females and 10 males (Osh-
ima et al., 1977, p. 100; Negayama et al., 1986,
p. 367; cf. Table 40); in the unsatisfactory sample
of two infant females, mean duration of the ges-
tation period is 170.5 days (extremes, 169.5-
171.5 days), and in the more satisfactory sample
of 10 infant males, the mean (±SD) duration is
175.4 ± 7.10 days (extremes, 161-184.5 days).
More data are required to ascertain whether the
mean gestation period of male infants exceeds
that of female infants.
FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES
63
Table 40. Gestation period duration in Macaca fuscata.
Publi-
Gestation
period
duration (days)
Refer-
cation
date
Mean
SD
Extremes
N
Basis of conception date estimate
ences-
1966
175.5-^
_
_
1
Estrous period
1
1976
173.0
6.9
161-188
17
48-hour controlled mating period
2
1977
176.9^
5.5
169.5-184.5
5
Copulatory period
3
1986
173.0^
7.4
161-179.5
V
Unspecified
4
1988
167.4
7.2
—
14
Ovulation'' (n = 6); 11 -day con-
trolled mating period (n = 8)
5
1990
162.8
3.6
157-180
6
5-day controlled mating period
6
1992
173.5
1.7
161-183
11
Ovulation^
7
1998
169.5
—
168-171
2
Artificial insemination
8
2000
166
—
—
1
In vitro fertilization''
9
2003
176.3
7.1
170-188.5
9
Ovulation'*
10
Totals
171.7
—
157-188.5
73
—
—
' Excludes two imprecise estimates: Tokuda (1961-62, p. 1 1) — 150-170 days; Hazama (1964, p. 89) — 5-6 months.
- Key to references: 7. Kawai (1966, p. 391). 2. Nigi (1976, p. 83). 3. Oshima et al. (1977, p. 100). 4. Negayama
et al. (1986, p. 367). 5. Shimizu (1988, p. 252). 6. Nozaki et al. (1990, p. 446). 7. Mitsunaga et al. (1992, p. 29). 8.
Torii and Nigi (1998, p. 401). 9. Torii et al. (2000, p. 44). W. Fujita (2003, p. 72).
' Mean based on midpoint(s) of duration range(s) specified in reference.
'' Excludes one stillbirth and one imprecise estimate (135-176 days).
-'' Determined by hormonal analysis.
^ Embryo subsequently implanted in fallopian tube.
During most pregnancies at Primate Research
Institute, Kyoto University, implantation bleeding
was detected for 3-7 days, beginning ca. 25-30
days after conception (Oshima, 1980, p. 4). At
Arashiyama West, Texas, females pregnant with
female fetuses reportedly gave and received more
aggression than females pregnant with male fe-
tuses, particularly during the first 2-3 months of
pregnancy (Noyes, 1981, p. 317).
Parturition
The time of parturition in M. fuscata is best
known, as would be expected, in caged popula-
tions (Itoigawa & Tanaka, 1963, p. 76; Oshima,
1980, p. 7; Negayama et al., p. 367; Kanazawa &
Nakamichi, 1991, p. 424). Of 16 reported partu-
ritions in caged females, 12 (including one or two
stillbirths) occurred near midnight during the in-
terval 2300-0300, three occurred during the in-
terval 2000-2200, and one occurred at 0727. In
one seminatural troop, a parturition occurred at
ca. 1800 (Fedigan & Fedigan, 1977, p. 205), and
in two natural troops, parturitions occurred at
1655 and ca. 0745 (Nakamichi et al., 1992, p.
414; Thomsen, 1997, p. 339). Other, less precise
reports concerning seminatural and natural troops
similarly indicate that parturition in these troops
usually occurs at night or during early morning
hours (Hazama, 1964, p. 83; Murray & Murdoch,
1977, p. 818; Pavelka, 1993, p. 61; Thomsen &
Solis, 2000, p. 690).
Shortly before parturition, prepartum females in
seminatural or natural troops apparently move
some distance from other troop members, fre-
quently into concealing underbrush (Fedigan &
Fedigan, 1977, p. 205; Nakamichi et al., 1992, p.
414; Fedigan & Zohar, 1997, p. 172; Thomsen,
1997, p. 339); all three or four reported parturi-
tions in seminatural or natural groups occurred on
the ground, not in trees. In three parturitions, con-
tractions were first observed 35, 44, and 51 min-
utes before birth of the infants (Itoigawa & Ta-
naka, 1963, p. 76; Nakamichi et al., 1992, p. 414;
cf. Kanazawa & Nakamichi, 1991, p. 424); in two
of these parturitions, rupture of the amniotic sac
reportedly occurred 4 minutes and 9 minutes be-
fore birth (Itoigawa & Tanaka, 1963, p. 76),
whereas in the third parturition, rupture of the sac
reportedly occurred 50 minutes before birth (O-
shima, 1980, p. 7). In caged, seminatural, and nat-
ural populations, the placenta apparently is almost
always completely consumed by the mother (Itoi-
gawa & Tanaka, 1963, p. 76; Hazama, 1964, p.
89; Fedigan & Fedigan, 1977, p. 206; Murray &
Murdoch, 1977, p. 818; Oshima, 1980, p. 7; Hirai-
wa, 1981, p. 315; Negayama etal., 1986, pp. 372-
373; Nakamichi et al., 1992, p. 414; Thomsen,
1997, p. 339; Thomsen & Soltis, 2000, p. 690).
64
FIELDIANA: ZOOLOGY
Table 41. Neonatal sex ratio in Macaca fuscata.
Number of births
Ratio
Sex
Observation
unspeci-
males/
Refer-
Locality
period
Females
Males
fied
Total
females
ences'
Arashiyama
1957-66
95
81
44
220
0.85
/
Arashiyama
1972-83
469
486
20
975
1.04
2
Awajishima
1978-96
281
311
—
592
1.11
3
Beaverton, U.S.A.
1966-75
-103
-74
—
-177
-0.72
4
Berlin, Germany
1977-95
55
58
4
117
1.05
5
Choshikei
1958-66
34
47
68
149
1.38
1
Funakoshiyama
1965-66
31
41
—
72
1.32
I
Gagyusan
1956-66
64
72
107
243
1.12
1
Hagachi
1966
5
2
2
9
0.40
1
Hakone-machi
1971-77
69
66
14
149
0.96
6
Houtosan
1963-66
33
28
—
61
0.85
1
Kanbanotaki
1962
15
12
—
27
0.80
I
Katsuyama-cho
1958-86
482
437
—
919
0.91
7
Kinkazan
1983-94
40
56
1
97
1.40
8
Kochi
1958-59
9
15
—
24
1.67
I
Kojima
1952-86
140
1602
21
321
1.14
9
Mihara-city
1964-65
5
9
—
14
1.80
I
Minoo-city
1957-66
117
148
—
265
1.26
1
Miyajima-cho
1962-66
20
25
6
51
1.25
1
Nametoko
1963-66
9
13
24
46
1.44
1
Ohirayama
1957-66
70
92
—
162
1.31
1
Okinoshima
1958-66
29
27
1
57
0.93
1
Rome, Italy
1977-88
37
48
—
85
1.30
10
Rosando
1958-62
36
44
—
80
1.22
1
Shiga'
1963-75
56
58
4
118
1.04
11
Taishakukyo
1958-62
19
20
—
39
1.05
1
Takagoyama (A, S)
1959-66
28
31
26
85
1.11
I
Takagoyama (I)
1974-78
53
36
—
89
0.68
12
Takaosan
1958-66
28
34
—
62
1.21
1
Takasakiyama
1956-62
260
289
118
667
1.11
13
Takasakiyama
1963-66
263
283
38
584
1.08
1
Takeno-ho
1965-66
7
14
4
25
2.00
1
Toimisaki
1958-65
40
43
22
105
1.08
1
Yaku.shima
1974-93
37
33
6
76
0.89
8
Totals
—
-3039
-3193
530
-6762
-1.05
—
' Key to references: /. Kawai et al. (1967, pp. 60-70). 2. Huffman (1991a, p. 33). 3. Nakamichi et al. (1997, p.
230). 4. Murray and Murdoch (1977, p. 818). 5. Harigel (1996b, p. 16). 6. Fukuda (1988, p. 481). 7. Itoigawa et al.
(1992, p. 57). 8. Takahata et al. (1988a, p. 341). 9. Watanabe et al. (1992, p. 8; cf. Iwamoto, 1974, p. 259). 10. Aureli
et al. (1990, p. 177). //. Tokita and Hara (1975, p. 27). 12. Hiraiwa (1981, p. 310). 13. Itani et al. (1964, p. 15).
- Includes one pair of male twins.
'' Shiga-A = Jigokudani.
Neonatal Sex Ratio; Birth Weight
In ca. 6232 M. fuscata births where sex of the
infant has been determined, the ratio of males to
females is ca. 1.05 (Table 41); this ratio is not
quite significantly different from 1.00 (x" = 3.806,
0.10 > f > 0.05). Neonatal sex ratios in M. fus-
cata apparently are not consistently correlated
with the mother's dominance rank (Wolfe, 1984b,
p. 141; Schino et al., 1999, p. 257; Brown & Silk,
2002, p. 11254).
Judging from the small samples of birth weight
that are available for M. fuscata (Table 42), mean
weight in female neonates (546.8 g, n = 11) ap-
pears to be fairly close to that in male neonates
(538.7 g, n - 10).
Twinning
In a translocated M. f. yakui population at Ohi-
rayama, one pair of twins was recorded among
FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES
65
Table 42. Birth weight (g) in Macaco fuscataJ
Females
Mean ± SD
Males
Sex unspecified
Mean ± SD
Mean ± SD
Refer-
ences-
600
536.75
600.0»
1
46
0
2
517 ± 90
4
—
0
546.8
11
500
Arashiyama
1
'450
Primate Research Institute, Kyoto University^
543.0 ±21.0^ 4 —
558.3' 3^ —
— 0 501.0 ± 52.9
460 1 —
Medical Lake Primate Field Station, U.S.A.
— 0 —
Tierpark Sababurg, Germany
580 1 —
Totals
538.7 10 -498.6
0
0
20
0
0
0
21
' Cf. Ohno et al. (1980, p. 936); Kurita et al. (2002, pp. 419-420).
- Key to references: 1. Hazama (1964, p. 91). 2. Oshima et al. (1977, p. 100). 3. Hamada (1982, p. 547). 4. Shimizu
(1988, p. 252). 5. Ota et al. (1991, p. 36). 6. Sirianni et al. (1975, p. 20). 7. Harigel (1998, p. 231).
' Supplementary information reported by Oshima (1980, p. 7): "'The birth weight of the offspring delivered naturally
ranged from 430 g to 610 g" (mean, sample size, and sex of infants not specified).
^Extremes, 518-570 g.
5 SD unspecified.
^ Age, in days, of infants when weighed: <1, 1, 5, 7.
^ Age, in days, of infants when weighed: <1, 1,6.
** Extremes, 545-655 g.
234 births that occurred during 13 reproductive
seasons (1957-1969); in a natural M. f. fuscata
population at Takasakiyama, one pair of twins
was recorded among 488 births that occurred dur-
ing 13 reproductive seasons (1977-1989); and in
another natural M. f. fuscata population at Koji-
ma, one pair of twins was recorded among 320
births that occurred during 35 reproductive sea-
sons (1952-1986) (Tanaka et al., 1970, p. 114;
Soumah & Yokota, 1992, p. 16; Watanabe et al.,
1992, p. 8); the composite twinning rate for these
three populations is 3/1042 = 0.29% (cf. Geiss-
mann, 1990, p. 391). Two additional pairs of
twins in two natural M. f. fuscata populations
(sample sizes not specified) have been reported by
Nakamichi (1983, p. 576), and one additional pair
of twins in a seminatural population (sample size
not specified) has been reported by Pavelka (1993,
p. 18).
At Arashiyama, a female nurtured a pair of "ar-
tificial twin" infants for at least 8 months (Ogawa,
1998, p. 101); this pair of infants consisted of one
male bom to the female on 13 May 1987 and a
second male neonate adopted by her 1 day later.
Birthrate
The mean annual birthrate (births/sexually ma-
ture females/year) in provisioned M. fuscata
troops (ca. 0.57) apparently exceeds that in non-
provisioned troops (ca. 0.38) (Table 43; see
above. Natural History: Group Size and Compo-
sition); corresponding mean interbirth intervals
are 1.75 years and 2.63 years, respectively (cf.
Ross, 1992, p. 209; Takahashi, 2002, p. 145). In
provisioned troops, variations in the level of pro-
visioning are positively correlated with variations
in the birthrate (Mori, 1979b, p. 375; Watanabe et
al., 1992, p. 7), and in nonprovisioned troops, an-
nual variations in fruiting rates during mating and
pregnancy seasons are positively correlated with
variations in birthrates during subsequent birth
seasons (Suzuki et al., 1998, p. 318). In the pro-
visioned natural group at Arashiyama, the mean
number of lifetime births per adult female report-
edly was approximately eight (maximum, 13)
(Koyama et al., 1975, p. 416).
In females whose infants have survived for at
least 1 year, intervals to the next birth are great-
66
FIELDIANA: ZOOLOGY
Table 43. Annual birthrate (births/sexually mature females/year) in Macaca fuscata.
Locality
Observation
period
Number
of births
Annual
birth rate
References'
Natural groups, nonprovisioned
Kawaradake
1971-78
80
0.59
/
Kinkazan
1983-92
249
0.36
2
Kojima-
1952-86
320
0.36
3
Ryozen
1974-80
45
0.34
4
Shimokita, SW
1965-74
10
0.31^
5
Yokoyugawa Valley''
1970-75
-11
-0.25
6
Yakushima (M. / \akui)
Kojiba^
1974-79
45
0.45
7
Mt. Kuniwari^
1975-78
27
0.46
7
Nine troops^
1989-95
46
0.28
2
Totals
Natural groups,
-833
provisioned
-0.38
—
Arashiyama
1954-83
973
0.62
8
Hakone-machi
1971-77
149
0.55
9
Katsuyama-cho
1958-86
919
0.44
10
Ryozen
1969-73
48
0.59
4
Shiga-A
1963-75
118
0.51
11
Takagoyama
1974-78*
87
0.53
12
Takasakiyama
1971-75
1406
0.62
6
Totals
—
3700
0.57
—
Seminaturai groups, provisioned
Aarashiyama West, U.S.A.
1972-79
99
0.51"
13
Beaverton, U.S.A.
1966-67
15
-0.79
14
Berlin, Germany
1977-95
117
0.46
15
PRIKU
1989-90
19
0.50
16
Miyajima-cho'°
1969
13
0.50
17
Rome, Italy
1977-81
46
0.86
18
Totals
—
210
-0.58
—
' Key to references: 1. Ikeda (1982, p. 344). 2. Suzuki et al. (1998, pp. 315-316; cf. Takahata et al., 1998b, p.
247; 1998a, p. 341); Sugiura et al. (2002, p. 76). 3. Watanabe et al. (1992, p. 8). 4. Sugiyama and Ohsawa (1982b,
p. 248). 5. Azuma (1985, pp. 3-4). 6. Yoshihiro (1985, p. 55). 7. Maruhashi (1982, pp. 321-322; cf. Azuma, 1985,
p. 2). 8. Koyama et al. (1992, p. 42). 9. Fukuda (1988, p. 481). 10. Itoigawa et al. (1992, p. 56). //. Suzuki et al.
(1975, p. 18). 12. Hasegawa and Hiraiwa (1980, p. 130). 13. Gouzoules and Goy (1983, p. 42). 14. Alexander and
Bowers (1967, p. 335). 15. Harigel (1996, p. 16). 16. Nozaki et al. (1995, p. 1251). 17. Stephenson (1975, p. 66).
18. Scucchi (1984, p. 204).
- Population variably provisioned.
^ 10 births/32 females.
■* Yokoyugawa Valley = Shiga-B (see Shiga-kogen).
"^ Includes daughter troops M, N, A, and H (products of successive fissioning); during the period 1984-89, the
annual birth rate of M troop decreased to 0.15 (Takahata et al., 1994, p. 318).
* Seven troops, excluding Kojiba troop.
7 Troops B, CC, G, H. M, NNA, P, S, T
* Takagoyama. provisioning discontinued April 1976.
■* Omitted from totals.
'° Translocated from Shodoshima in 1962.
er — and birthrates are correspondingly lower —
than in females whose infants have not survived
(Table 44; cf. Tanaka et al., 1970. p. 115; Nomura
et al., 1972, p. 478; Mitsunaga et al., 1992, p. 25;
Takahashi, 2002, p. 145). Lactation apparently
tends to suppress the ovarian endocrine function
of females (Hiraiwa, 1981, p. 312; Yoshida et al.,
2001, p. 371).
The age-specific birthrate in M. fuscata appar-
ently tends to increase during the first few repro-
ductive years, tends to maintain a relatively high
level during the middle reproductive years, and
probably tends to decline during the last few re-
productive years (Fig. 23; cf. Wolfe & Noyes,
1981, p. 699; Gouzoules et al., 1982, p. 1140;
Sugiyama & Ohsawa, 1982b, p. 246; Fukuda,
FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES
67
Table 44. Effect of Macaca fuscata infant death on
interbirth interval of mother.
Mean interbirth
interval (yr)
Locality
Following
survival
of infant
to age 1 yr
Following
death of
infant
before age
1 yr
Refer-
ences'
Natural groups, nonprovisioned
Kinkazan 2.37 1 .59
Yakushima 2.24 1.50
Natural groups, provisioned
Arashiyama i.46 1.15
Katsuyama-cho 1.58 1.29
Seminatural group, provisioned
Rome 1.22- 1.06^
' Key to references: /. Takahata et al. (1998a, p. 341).
2. Scucchi (1984, p. 205).
^ Survival of infant to age 6 months.
^ Death of infant before age 6 months; includes still-
births.
1988, p. 482; Watanabe et al., 1992, p. 17; Hari-
gel, 1996, p. 16; Shimizu, 1998, p. 1 15). In seven
groups where the relationship between birthrate
and female dominance rank has been studied (Ta-
ble 45), birthrate is positively correlated with
dominance rank in two groups, birthrate is not
correlated with female dominance rank in four
groups, and evidence is equivocal in one group.
Infant Mortality
The infant mortality rate (deaths before age 1
year/births) in nonprovisioned groups of M. fus-
cata (mean = 28.4%) apparently is more than
twice as great as that in most provisioned natural
groups (9.4%) and seminatural groups (8.5%) (Ta-
ble 46); the high infant mortality rate reported in
the Hakone T provisioned natural group (49.7%)
is exceptional and remains unexplained (Fukuda,
1988, p. 483). Approximately half of all infant
deaths probably occur before age 1 month (Itoi-
gawa et al., 1992, p. 63; Koyama et al., 1992, p.
41 ; Watanabe et al., 1992, p. 22; Nakamichi et al.,
1997, p. 233).
Known violent causes of infant deaths include
attacks by dogs or raccoon dogs (Iwamoto, 1974,
p. 257) and attacks by adult male monkeys (Kur-
land, 1977, p. 83; Soltis et al., 2000, p. 197; Taka-
hashi, 2002, p. 151). Mothers have been observed
to carry the bodies of stillbirths or dead infants
for several days, even after the bodies have begun
to decompose (Green, 1975, p. 15; Nakamichi et
al., 1983, p. 64; Pavelka, 1993, p. 63; Takahashi,
2002, p. 144).
Nursing; Weaning
Three observed newborn M. fuscata infants be-
gan to suckle 23 minutes, ca. 90 minutes, and 1 10
minutes after parturition (Itoigawa & Tanaka,
1963, p. 76; Thomsen, 1997, p. 339). During the
Table 45. Relationship between birthrate and female dominance rank in nonprovisioned and provisioned groups
of Macaca fuscata.
Locality
Birthrate significantly
Observation Number correlated with female
period of births dominance rank?
References'
Kinkazan
Kojima-
Yakushima
Arashiyama
Arashiyama West""
Katsuyama-cho
Takasakiyama
Nonprovisioned groups
1983-94
97
No
1
1952-86
258
Evidence equivocal
2
1974-93
76
Provisioned groups
No
1
1954-83
975
No
3
1976-78
>82
No
4
1958-84
919
Yes
5
1977-89
488
Yes
6
' Key to references: /. Takahata et al. (1998a, p. 345). 2. Mori (1979b, p. 388); Watanabe et al. (1992, p. 11). 3.
Wolfe (1984b, p. 139); Koyama et al. (1992, p. 42); cf. Fedigan et al. (1986, p. 151). 4. Gouzoules et al. (1982, p.
1141); Fedigan et al. (1986, p. 151). 5. Itoigawa et al. (1992, p. 65). 6. Soumah and Yokota (1992, p. 15).
- Population variably provisioned.
^ South Texas, U.S.A.
68
FIELDIANA: ZOOLOGY
Table 46. Infant mortality rate (infant deaths before age 1 yr/births) in nonprovisioned and provisioned groups
of Macaca fuscata. '
Locality
Observation
years
Number
of births
Infant mortality
rate (%)
References^
Kinkazan
Kojima'
Shiga B-2
Shimoicita SW
Yakushima
Totals
Arashiyama
Awajishima
Hakone T
Katsuyama-cho
Shiga A
Takagoyama-I
Takasakiyama
Totals
Arashiyama West''
Natural groups, nonprovisioned
1983-94
97
22.7
1
1952-86
320
31.2
2
1970-75
15
40.0^
3
1965-74
9
0
4
1974-93
76
25.0
1,4,5
—
517
28.4
—
Natural
groups, provisioned
1954-83
953
10.3
6
1978-95
543
12.3
7
1971-77
149
49.7
8
1958-85
949
10.2
9
1963-75
118
6.85
3, 10
1975-78
68
16.2
11
1956-62
667
4.5
12
1987-89
215
882
8.8
5.6
13
—
3513*
9.46
—
Seminatural group, provisioned
1972-93 1230
8.5
14
' For mortality rate to age 2 years at Ryozenyama, see Sugiyama and Ohsawa (1982b, p. 250) and Takahata et al.
(1998a, p. 341).
- Key to references: 1. Takahata et al. (1998a, p. 341). 2. Mori (1979b, p. 375); Watanabe et al. (1992, p. 8). 3.
Suzuki et al. (1975, pp. 19, 21; cf. Yoshihiro, 1985, p. 55). 4. Azuma (1985, pp. 3, 4). 5. Maruhashi (1982, p. 322).
6. Koyama et al. (1992, p. 41). 7. Nakamichi et al. (1997, p. 233). 8. Fukuda (1988, p. 481). 9. Itoigawa et al. (1992,
p. 63). 10. Tokita and Hara (1975, p. 27). //. Hiraiwa (1981, p. 319). 12. Itani et al. (1964, p. 15). 13. Soumah and
Yokota (1992, p. 17). 14. Fedigan and Zohar (1997, p. 168).
^ Population variably provisioned.
^ Excludes two miscarriages and/or stillbirths.
■* Excludes seven miscarriages and/or stillbirths.
*> Excludes aberrant Hakone T data.
^ South Texas, U.S.A.
first 2 weeks of life, 33 infants in a seminatural
group spent an average of 87% of observed in-
tervals in contact with their mothers' ventrum;
69% of these intervals included infants' oral con-
tact with their mothers' nipples (Murray & Mur-
doch, 1977, p. 819). Infants reportedly begin to
taste solid food at age 2 weeks, and while con-
tinuing to nurse, they begin to eat solid food at
age 4-6 weeks (Pocock, 1906, p. 567; Hasegawa
& Hiraiwa, 1980, p. 135; Hiraiwa, 1981, p. 313).
When an infant is approximately 6 months old —
which is near the beginning of the troop's next
mating season — its mother's milk secretion
abruptly decreases (Ota et al., 1991, pp. 41, 47;
Tanaka, 1992, p. 133), and when an infant is 7-
10 months old, its mother may reject the infant's
attempts to suckle (Fedigan & Fedigan, 1977, p.
217; Hasegawa & Hiraiwa, 1980, p. 135; Hiraiwa,
1981, p. 318); such rejection reportedly is more
frequent in estrous females than in nonestrous fe-
males (CoUinge, 1987, p. 142; 1991, p. 163; Pav-
elka, 1993, p. 66). Although some infants may be
fully weaned by age 6-8 months (Pocock, 1906,
p. 567; Tanaka et al., 1970, p. 116; Ota et al.,
1991, p. 47; Pavelka, 1993, p. 66), some continue
to nurse to age ca. 1 year — until just before their
mother's next parturition (Tanaka, 1992, p. 136;
Inoue et al., 1993, p. 505), and others may con-
tinue to nurse to age ca. 2.5 years — if their moth-
ers have had no intervening parturitions (Fedigan
& Fedigan, 1977, p. 217; Hiraiwa, 1981, p. 315;
Sugiyama & Ohsawa, 1982b, p. 250; Nakamichi,
1989, p. 739; Tanaka, 1992, p. 136; Pavelka,
1993, p. 66; Tanaka et a!., 1993, p. 171).
Adult males oi M. fuscata, including high-rank-
ing males, occasionally have been observed to
FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES
69
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70
FIELDIANA: ZOOLOGY
Table 47. Mean life span in Macaca fuscata females.
Locality
Number of
Birth period of deceased
females studied females
Mean (±SD)
female life span
(yr)
References'
Yakushima: M. X P
Natural population, nonprovisioned
ca. 1974-93 36
6.3 ± 5.2
/
Takasakiyama
Natural population, provisioned
ca. 1952-73 —
8.4 ± ?
2
Arashiyama AAVest
Natural/seminatural- population, provisioned
1950-67 72 13.6 ± 9.4^
3
Key to references: /. Takahata et al. (1998a, p. 342). 2. Masui et al. (1975, p. 405). 3. Fedigan (1991b, p. 142).
Natural provisioned population prior to February 1972; seminatural provisioned population subsequently.
For this calculation, reported deaths at unspecified ages < 1 year were arbitrarily set at 0.5 years.
protect and groom infants (Itani, 1959, p. 89; Ta-
kahata, 1982a, p. 14; Yoshimura, 1984, p. 25).
Longevity; Reproductive Senescence
The greatest accurately known life spans in M.
fuscata are 33 years in a female in a provisioned
natural group (Koyama et al., 1992, p. 37; of. Pav-
elka & Fedigan, 1999, p. 458; Fedigan & Pavelka,
2001, p. 113) and 28 years in a male in another
provisioned natural group (Nakamichi et al.,
1995, p. 386); greater estimated ages of 30-40
years have been reported for four males, but the
actual birth dates of these males are unknown
(Itani et al., 1964, p. 18; Tasumi, 1969, p. 263;
Itoigawa, 1982, p. 380). Mean female life spans
apparently are much less than half of the greatest
known female life span; reported mean female life
spans are 6.3 years in a nonprovisioned natural
population, 8.4 years in a provisioned natural pop-
ulation, and 13.6 years in a provisioned natural/
seminatural population (Table 47). Because males
usually emigrate from their natal troops, deter-
mination of mean male life spans in unconfined
populations is impractical; in a confined popula-
tion, the mean male life span apparently was ca.
60% of the mean female life span (Fedigan &
Zohar, 1997, p. 165).
The greatest age at which a female M. fuscata
is known to have borne an infant is 26 years (Itoi-
gawa et al., 1992, p. 56; Shimizu, 1998, p. 115);
this occurred in a provisioned natural group at
Katsuyama-cho. In a provisoned natural group at
Arashiyama, the mean (±SD) age at last parturi-
tion in seven females that survived to age 20 years
was 21.7 ± 2.0 years (extremes, 19-25 years)
(Takahata et al., 1995, p. 172); similarly, at Ara-
shiyama West, South Texas, the mean age at last
parturition in 16 females that survived to age 20
years was 21.3 ± 3.2 years (extremes, 11-25
years) (Pavelka & Fedigan, 1999, p. 458). The
mean postreproductive interval between a fe-
male's last parturition and her disappearance
(probable death) from the Arashiyama group was
6.0 ±3.5 years (1-10 years, n = 8) (Takahata et
al., 1995, p. 172), and the corresponding mean
postreproductive interval between a female's last
parturition and her death in the Arashiyama West
group was 5.0 ± 2.5 years (2-11 years, n = 20)
(Pavelka & Fedigan, 1999, p. 458). Postreprod-
uctive females may contribute to the fertility of
their daughters, and they probably contribute to
the survival of their daughters' offspring (Pavelka
et al., 2002, p. 412).
Although the sexual activity of females appar-
ently tends to decline during the postreproductive
interval, females at this stage often continue to
have estrous cycles and to engage in copulation
(Koyama et al., 1992, p. 37; Mitsunaga et al..
Fig. 23. Age-specific birthrates in two nonprovisioned troops (solid symbols) and two provisioned troops (open
symbols) oi Macaca fuscata: nonprovisioned troops — Kinkazan, 1983-1994, 97 births, and Yakushima, 1974-1993,
76 births (Takahata et al., 1998, pp. 343-344); provisioned troops — Arashiyama, 1954-1983, 975 births (Koyama et
a!., 1992, p. 37); Katsuyama-cho, 1958-1986, 919 births (Itoigawa et al., 1992, p. 56).
FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES
71
1992. p. 28: Takahata et al.. 1995. p. 174): cop-
ulation by a 32-year-old female has been reported
(Oda & Masataka. 1995. p. 135). The ovaries of
postreproductive females reportedh are reduced
in size, impaired histologically, and impaired en-
docrinologically (Mitsunaga et al.. 1992. p. 26:
Nozaki et^al.. 1995. p. 1253: 1997. p. 92: Shimizu.
1998. pp. 115. 117: Yoshida et al.. 2001. p. 370).
A male M. fitscata at KatsuN ama-cho \\ as ob-
served to copulate at age ca. 27 years, and another
male at Takasakiyama was observed to chase an
estrous female at age ca. 30 years (Itoiga\\a.
1982. p. 382). At Arashiyama West, however, a
male of estimated age >20 years rarely main-
tained an erection, although he consorted with five
estrous females. The testes and accessory glands
of males of estimated age ^26 % ears \\ ere found
to be degenerate relati\e to those of males age
20-2 1 \ ears, although a few sperm \\ ere noted in
the seminiferous tubules of males near age 30
years (Matsubayashi et al.. 1994. pp. 201-204).
Population Growth Rate
Predictably, the mean population growth rate in
nonprovisioned natural M. fuscoto groups (ca.
2.5^^) is considerably less than that in provisioned
natural groups (ca. 9.0*^) and in provisioned
seminatural groups (ca. 12.3^) (Table 48).
Fossils
Fossils referred to Macaco Jiiscata. associated
w ith numerous other fossil mammals, have been
unearthed at various localities on Honshu. Shi-
koku and Kyushu (Tables 49. 50: Fig. 24). Based
on tephrostratigraphy and oxygen isotope strati-
graphy of Japanese Pleistocene proboscidean spe-
cies, it has been concluded that dry-land connec-
tions ("land bridges") between the Japanese is-
lands and the adjacent Asian continent existed
twice during the Middle Pleistocene (Kawamura.
1998. p. 252: Dobson & Kawamura. 1998. p. 387:
Konishi and Yoshikawa, 1999. p. 131). These land
bridges apparently occurred at oxygen isotope
stage 16 (0.63 million years ago: Ma) and at ox-
ygen isotope stage 12 (0.43 Ma), when Stegodon
ohentalis and Palaeoloxodon naunianni. resp>ec-
tively, immigrated into the Japanese islands. S. or-
ientalis survi\ed for some time but apparently
was ultimately replaced by P. naunianni: these
two proboscideans were not coexistent.
To date, the purported oldest macaque fossil in
Japan is an isolated left lower third molar (mesio-
distal length. 12.1 mm: buccolingual width. 7.0
mm), comparable in size and shaf)e to correspond-
ing molars of the extant Japanese macaque, that
was obtained from a fissure deposit at Ando Quar-
ry. Yamaguchi Prefecture (Iwamoto and Hasega-
wa. 1972. p. 78: Aimi. 2002. p. 242). The asso-
ciated fossil proboscideans are S. orientalis and
P. naunianni, which indicates that the fossil as-
semblage at Ando Quarry is a mixed one. The age
of the fossil macaque molar remains elusive: if it
was originally associated with 5. orientalis, its age
is ca. 0.63 Ma. but if it was originally associated
with P. naunianni. its age is ca. 0.43 Ma.
Another early macaque fossil in Japan is a right
humerus, similar to that in extant male Japanese
macaques, that was unearthed at Yarimizu, Jizodo
Formation or Kamiizumi-Kiyokawa Formation,
Chiba Prefecture (Iwamoto and Hasegawa. 1991,
p. 100). The associated proboscidean fossils have
been identified as P. naunianni.
A\ailable evidence suggests that the ancestors
of M. fuscata dispersed from mainland Asia to the
Japanese Islands either ca. 0.63 Ma or ca. 0.43
Ma. No evidence exists concerning the possibility
of disp>ersal of M. fuscata ancestors to Japan be-
fore 0.63 Ma.
Several macaque fossils ha\e been obtained
from late Middle Pleistocene to Late Pleistocene
deposits on the Korean Peninsula, where no ma-
caques occur at present (Lee. 1983, p. 40: Park
and Lee, 1998, p. 56). Although some of these
Korean fossils haN e been referred to Macaca cf.
robitsta based on their molar size (Table 51), the
measurements of these fossil molars actually are
within the range of variation show n by Japanese
macaques (Table 52). Further comparative studies
of the Korean fossils and extant Japanese ma-
caques are required to in\estigate the possibility
of an evolutionary relationship.
Systematics
Geographic Variation and Subspecific
Recognition
Based on geographic variation of pelage color
and body size. Kuroda (1940, p. 273) proposed
that the population of M. fuscata that inhabits
72
FIELDIANA: ZOOLOGY
Table 48.
Population growth
rate in Macac
a fuscata
groups.
Number
Annualized
of
population
groups Census
Initial
Final
growth rate'
Refer-
Locality
observed interval
census
census
(%)
ences-
Natural groups, i
nonprovisioned
Kinkazan
5
1983-93
-270
-260
— 0.3^
1
Kojima^
1
1952-86
21
-100
-4.7
2
Shimokita M
1
1971-85
-24
-79
-8.9
3
Shimokita O
1
1963-81
-12
-20
-2.9
4
Takagoyama I'
1
1976-78
107
100
-3.3
5
Yakushima''
1-4
1974-90
42
-60
-2.2
6
Mean ± SD
—
Natural groups
i, provisioned
—
-2.5 ± 4.2
—
Arashiyama A and B
2
1954-72
-28
301
-14.1
7
Arashiyama B
1975-78
185
253
11.0
8
Awajishima
1978-95
-100
>250
-5.5
9
Gagyusan
1955-58
110
160
13.2
10
Hagachizaki
1955-59
>70
>100
-9.3
U
Hakone T
1956-67^
-35
-75
-7.2
12
Katsuyama-cho
1-2
1958-86
112
236
2.7
13
Shiga A
1963-75
23
95
12.5
14
Shiga A-l
1990-97
108
252
12.9
15
Takagoyama I'
1974-76
98
107
4.5
5
Takasakiyama
1-3
1951-92
166
1963
6.2
16
Mean ± SD
—
—
—
—
-9.0 ± 4.0
—
Seminatural groups, provisioned
Arashiyama West, U.S.A.
1972-94
150
>600
>6.5
17
Beaverton, U.S.A.
1964-78
46
316
14.8
18
Berlin, Germany
1977-95
9
77
12.7
19
Arnhem. Netherlands
1973-85
8
26
10.3
20
Montreal, Canada
1984-93
14
40
12.4
21
Nojima
1969-71
51
113
48.9«
22
Rome. Italy
1977-81
27
51
17.2
23
Mean ± SD
—
—
—
—
-12.3 ± 3.7"
—
' Annualized growth rate = [(C^/C,)"^] - 1, where Y = years in census interval, C, = initial census, and C2 -
final census.
- Key to references: 1. Takahata et al. (1998a, p. 340). 2. Watanabe et al. (1992, p. 4). 3. Watanuki et al. (1994, p.
16). 4. Izawa (1982. p. 13); Azuma (1985, p. 4). 5. Hasegawa and Hiraiwa (1980, p. 130). 6. Takahata et al. (1994,
p. 319; cf. Maruhashi, 1982, p. 330). 7. Koyama et al. (1975, p. 412). 8. Grewal (1980, p. 331; cf. Takahata, 1980,
p. 304). 9. Nakamichi et al. (1997, p. 226). 10. Furuya (1960, p. 149). //. Nishida (1966, p. 151). 12. Fukuda (1988,
pp. 480, 481). 13. Itoigawa et al. (1992, pp. 52, 53). 14. Suzuki et al. (1975, p. 18; cf. Tokita & Hara, 1975, p. 27).
15. Tanaka (1998. p. 1230). 16. Ohsawa and Sugiyama (1996. pp. 163, 167). 17. Jack and Pavelka (1997, p. 370).
18. Eaton (1976, p. 105); Rostal et al. (1986. p. 453). 19. Harigel (1996. p. 16). 20. Scheurer and Thierry (1985, p.
491). 21. Prud'homme and Chapais (1996, p. 432). 22. Nomura et al. (1972, p. 481). 23. Scucchi (1984, p. 204).
^ During the winter of 1983-84, an unusually heavy snowfall at this locality resulted in the deaths of ca. 90 monkeys
(Takahata et al., 1998a, p. 340).
■* Variably provisioned.
'' Provisioning of this group was discontinued in March 1976.
^ Kojiba troop and its daughter troops.
^ Excludes period after 1 967, when uncensused daughter troops separated from the main troop.
•* Birthrates in this group were artificially enhanced by removing infants from their mothers 3-6 months after birth.
' Excludes Nojima mean (see footnote 8).
Yakushima, a small island at the southern limit of
distribution of the species, should be recognized
as a distinct subspecies, M. f. yakui. Comparing
the Yakushima population with more northern
populations, Kuroda noted that pelage color is
darker — particularly on the back, limbs, and ex-
tremities— in the Yakushima population and that
body size is smaller in this population. Although
FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES
73
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74
FIELDIANA: ZOOLOGY
Table 50. Mammal stages, biozones, and stratigraphy of Quaternary deposits in Japan (Kamei et al., 1988;
Konishi & Yoshikawa, 1999).
Mammal stage
Biozone
Stratigraphy'
Oxygen isotope stage
Shitamachian
QMS
Hanaizumian
QM7
Upper Kuzuuan
QM6
QMS
Lower Kuzuuan
QM4
Katadaan
QM3
QM2
Akashian
QMl
Holocene
Uppermost Pleistocene
Upper Pleistocene, Lower
Middle Pleistocene, Upper
Middle Pleistocene, Middle
Middle Pleistocene, Lower
Lower Pleistocene, Upper
Lower Pleistocene, Lower
Stage 12 (0.43 Ma)
Stage 16 (0.63 Ma)
Cited as in original source.
overlap in the variation in pelage color and body
size in Yakushima and non-Yakushima popula-
tions probably is greater than implied by Kuroda,
one aspect of pelage color differentiation in the
Yakushima population appears adequate to war-
rant continued subspecific recognition of M. f.
yakui.
Pelage Color — Dorsal pelage color is relative-
ly dark in Yakushima M. fuscata and tends to be-
come paler in populations that inhabit increasing-
ly higher latitudes (Table 2); this pattern of geo-
graphic variation apparently applies to pelage col-
or in neonates as well as to that in adults
(Imaizumi, 1970, p. 285; Kuroda, 1984, p. 14; Ha-
mada et al., 1992, p. 7). However, dorsal pelage
in populations on two southern islands — Kojima
and Kyushu — apparently averages as dark as that
in the Yakushima population, at least in adults
(Table 2). The similarity of dark dorsal pelage col-
or in Yakushima and Shikoku specimens was pre-
viously noted by Thomas (1906 ["1905"], p. 361)
and Kuroda (1940, p. 273).
In Yakushima M. fuscata, pelage on the dorsal
surface of the hands averages distinctly darker
(blackish) than pelage on the dorsal surface of the
trunk, whereas in all non-Yakushima samples
studied — including Kojima and Kyushu sam-
ples— dorsal pelage on the hands does not average
darker than dorsal pelage on the trunk (Table 2).
Assuming that this difference is confirmed by the
evidence of future samples, particularly from
southern islands, this character state may be taken
as diagnostic of the Yakushima population of M.
fuscata; similar pelage characters have been used
to define the subspecies M. fascicularis atriceps
and M. f condorensis (Fooden, 1995, p. 68).
Body Size — Judging from available measure-
ments of sitting height and anterior trunk length,
body size in M. fuscata generally tends to increase
clinally with latitude from Yakushima to Honshu
(Tables 6, 7; Figs. 5, 6). Although body size in
Yakushima M. fuscata, at the southern extreme of
this cline, averages less than in non-Yakushima
M. fuscata, variation in body size in the Yakushi-
ma population apparently overlaps strongly with
that in non-Yakushima populations; this overlap-
ping variation undermines the usefulness of body
size in subspecific definition within M. fuscata.
Patterns of variation in body weight and greatest
length of skull are generally similar to those in
sitting height and anterior trunk length (Tables 1 1 ,
19; Figs. 10, 15), which further indicates that
body size probably is unsuitable for defining sub-
species in this species.
Cranial Morphology — As indicated above,
greatest length of skull varies clinally in M. fus-
cata (Table 19; Fig. 15). Although greatest length
of skull averages less in Yakushima M. fuscata
than in non-Yakushima M. fuscata, variations of
these two populations strongly overlap.
The orbits in Yakushima M. fuscata average
narrower than in non-Yakushima M. fuscata (Ta-
ble 21; Fig. 17; cf. Ikeda & Watanabe, 1966, p.
272; Kuroda, 1984, p. 15). In these populations,
the overlap in orbital breadth measurements is rel-
atively small (1.4 mm in females, 1.4 mm in
males).
Mitochondrial DNA; Blood Proteins —
Available evidence concerning geographic varia-
tion in mtDNA and blood proteins fails to indi-
cates clear-cut differences between Yakushima M.
fuscata and non-Yakushima M. fuscata (Figs. 1 8-
20). Mitochondrial DNA in the Gagyusan (Hon-
shu) sample is more similar to that in the Yaku-
shima sample than to that in other Honshu sam-
ples, and blood proteins in the Boso Peninsula
(Honshu) sample likewise are more similar to
FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES
75
Yarimizu
Ojikado
'Bimemi
7
Tengakuin
Gansuiji
shikawa Shiraiwa
Suse Yage
Fig. 24. Distribution of fossil macaque localities in Japan: for details, see Table 49.
76
FIELDIANA: ZOOLOGY
Table 51. Dental measurements (mm) in four Korean fossil macaque specimens collected at Turubong Cave
(Lee, 1983, p. 40) and one Japanese fossil macaque specimen collected at Ando Quarry (Iwamoto & Hasegawa,
1972, p. 79).
Korean fossils
. Japanese fossil
mandibular
Maxillary
teeth
Mandibular teeth
No.
2TB-947.'
No.
2T-322,
tooth
Dental dimension
6
(Unknown)
6 No.
2T-1488
(Unknown)
P3-M3 length
_
43.4
M1-M3 length
29.5
—
29.7
—
—
M2-M3 length
—
23.2
—
—
—
P3 length
—
—
6.4
—
—
P3 breadth
—
—
4.8
—
—
P4 length
—
—
6.4
—
—
P4 breadth
—
—
6.2
—
—
Ml length
8.0
—
7.9
—
—
M 1 greatest breadth
7.6
—
7.1
—
—
M2 length
10.0
—
9.4
9.6
—
M2 greatest breadth
9.1
—
8.0
8.2
—
M3 length
10.5
—
12.6
13.0
12.1-
M3 greatest breadth
9.0
7.8^
7.6
7.8
7.0-
' Referred by Lee (1983) to Macaca robusta; cf. Fooden, 1990, p. 656.
- Left mandibular M3.
^ Mandible height at M3 = 23.2 mm.
those in the Yakushima sample than to those in
other Honshu samples.
Key to External Characters of
Recognized Subspecies
Pelage on dorsal surface of hands blackish, dis-
tinctly darker than pelage on dorsal surface of
trunk yakui
Pelage on dorsal surface of hands pale brown to
dark brown, color similar to or paler than that
of pelage on dorsal surface of trunk . . fuscata
Japanese Vernacular Names Applied to
Macaca fuscata
Kishida (1953, p. 3) has published the follow-
ing list of vernacular names that have been ap-
plied to the Japanese macaque: awasaru, chichi-
busaru, enko, enkou, ete, etekou, etemono, hon-
saru, hondozaru, isonotamotomai, isonotachihaki,
kimura, koganomiko, kokawasaru, kokesaru,
koko, konohasaru, konomitori, makkou, mame-
saru, masaru, mashi, mashiko, mashira, mashita,
miko, mikou, minosaru, mokkaku, mokko, mok-
kou, mouzoku, nihonsaru, nihonzaru, nipponsaru,
oise, sakebi, sakebitori, sakebu, saru, sarumaru,
sarumaro, sarumatsu, sikaisaru, shikokusaru, su-
zunomiko, taka, takanomiko, tosasaru, toshuusa-
ru, yaen, yaenbo, yakushimazaru, yamasaru, yo-
buko, yobukotori.
Subspecies Accounts
Macaca fuscata fuscata (Gray, 1870)
Synonymy
Monkeys of a docile kind, with short tails:
Kaempfer, 1727, p. 126 — external characters.
[M]onkeys of a small race: Golovnin, 1824, p.
190 — geographic distribution, "southern and
middle provinces of the empire."
Simla (Cercopithecus) japonica: C. G. C. Rein-
wardt, 24 February 1821, unpublished letter in
RMNH archives — unavailable name, applied to
skeleton shipped to RMNH.
[PJapio Japonicus [Ogilby], 1838, pp. 364 —
name applied to specimens "recently brought
from Japan by Dr. S[ie]bold." Anderson, 1879,
p. 78 — footnote citation. Fooden, 1966, p.
159 — proposed for suppression as neglected
name. Fooden, 1967, p. 250 — formal request
for suppression of name by International Com-
mission on Zoological Nomenclature. Interna-
FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES
77
Table 52. Dental measurements (mm) in samples of extant Macaca fuscata (reference: Kondo, 1 987, p. 29), for
comparison with corresponding measurements in Korean and Japanese fossil macaques (see Table 51).
Tooth
Sex
N
Mean
SD
Min.
Max.
N
Mean
SD
Min.
Max.
Max:
illary teeth:
length
Maxillary
teeth: greatest breadth
Ml
M
84
7.87
0.366
7.00
8.70
84
7.68
0.401
6.80
8.65
F
72
7.54
0.379
6.35
8.45
72
7.38
0.332
6.70
8.35
M2
M
92
9.26
0.424
8.15
10.30
88
9.27
0.375
8.40
10.25
F
85
8.93
0.470
7.65
10.15
84
8.75
0.417
7.65
9.90
M3
M
76
9.39
0.437
8.30
10.70
74
9.12
0.338
8.30
9.75
F
68
8.90
0.536
7.85
10.15
61
8.58
0.370
7.65
9.40
Mandibular teeth:
length
Mandibular
■ teeth: greatest breadth
P3
M
86
6.62
0.542
5.65
7.75
92
5.08
0.459
4.15
6.10
F
68
5.49
0.282
4.95
6.15
76
4.56
0.288
3.90
5.30
P4
M
78
6.05
0.280
5.30
6.55
81
5.37
0.310
4.70
6.10
F
67
5.68
0.290
5.00
6.40
69
5.13
0.344
4.25
5.95
Ml
M
74
7.83
0.289
7.20
8.55
70
5.92
0.304
5.30
6.60
F
62
7.50
0.364
6.70
8.30
61
5.73
0.342
4.95
6.55
M2
M
85
9.12
0.367
8.20
10.05
85
7.32
0.383
6.55
8.40
F
67
8.64
0.357
7.75
9.40
72
6.92
0.385
6.10
7.65
M3
M
63
11.60
0.608
10.30
13.25
69
7.67
0.345
7.00
8.60
F
57
11.02
0.601
9.35
12.45
61
7.21
0.474
5.85
8.35
tional Commission on Zoological Nomencla-
ture, 1970, p. 77 — name suppressed.
Japansche aap: C. G. C. Reinwardt, 12 May 1822,
unpublished letter in RMNH archives — applied
to skeleton shipped to RMNH.
Speciosus: F. Cuvier, 1825, livraison 47, p. 2 —
unavailable specific name, not published in
combination with generic name; based on col-
ored illustration of "Macaque a face rouge"
(unnumbered plate), received from A. Duvaucel
and R Diard; habitat mistakenly given as "les
Indes orientales."
Macacus speciosus I. Geoffroy, 1826, p. 589 —
binomial based on Speciosus: F. Cuvier, 1825.
Lesson, [1830], p. 121 — habitat mistakenly giv-
en as Sumatra. I. Geoffroy, 1831, p. 63 — spe-
cies recorded as doubtfully distinct. Sclater,
1860, p. 420 — external characters; taxonomic
comparisons. Sclater, 1865 ("1864"), pp. 709,
710 — captive reported in London. Anderson,
1875 ("1874"), p. 652 — taxonomic relation-
ships. [Sclater], 1875 ("1874"), p. 685— cap-
tive imported from Japan. Sclater, 1875, p. 418,
plate 47 — captive obtained in Kyoto; external
characters. BIyth, 1875, p. 6 — name said to be
wrongly applied to Japanese macaque (mistak-
enly applied by Blyth to Indochinese stump-
tailed macaque). Schlegel, 1876 — type series
cataloged. Sclater, 1876, p. 332 — name rejected
as possibly based on misidentified holotype.
Anderson, 1879, p. 50 — taxonomic history;
name mistakenly regarded as applicable to In-
dochinese stumptailed macaque. Jentink, 1887,
p. 27 — type series cataloged. Jentink, 1892, p.
31 — type series cataloged. Sclater, 1892, p.
471 — name accepted as correctly applicable to
Japanese macaque. Forbes, 1894, p. 8 — name
erroneously included in synonymy of Indochi-
nese stumptailed macaque. Sclater and Sclater,
1899, p. 227 — zoogeography. Fooden, 1967, p.
250 — formal request for suppression of name
by International Commission on Zoological No-
menclature. International Commission on Zoo-
logical Nomenclature, 1970, p. 77 — name sup-
pressed.
Macaca speciosa: Fooden, 1966, p. 153 (not Po-
cock, 1926 ["1925"], pp. 1497, 1535— mis-
identification) — spelling of generic name cor-
rected, gender ending of specific name changed
accordingly; taxonomic history; suppression of
name proposed to avoid bibliographic confu-
sion.
S[imia] speciosa: Fischer, 1829, p. 30 — new com-
bination; habitat, "In India orientali." de Blain-
ville, 1838, p. 361 — habitat, Japan.
Inuus speciosus: Temminck, 1836, p. xxii — new
combination; taxonomic history; habitat, Japan.
Temminck, 1838-1839, p. 281— compared with
M. syhanus. Temminck, 1842, p. 9, plates 1,
2 — taxonomic history; geographic distribution;
natural history. Von Siebold, 1852, p. 245 —
name cited from Temminck, 1842. Rein, 1875,
p. 55 — geographic distribution extends north-
ward to 4rN.
78
FIELDIANA: ZOOLOGY
Innuus speciosus: Temminck, 1842, plates I, II —
misspelling of generic name.
Papio speciosus: Ogilby, 1839, p. 49 — new com-
bination; taxonomic history; habitat, Japan.
Cynopithecus speciosus: Lesson, 1840, p. 102 —
new combination; habitat, Japan. I. Geoffroy,
1851, p. 32 — species reassigned to Macacus.
Pith[ecus] (Macfacus]) speciosus: Dahlbom,
1856, p. 116 — new combination.
Affenspezies: P. F. von Siebold, 12 February 1829,
unpublished letter in RMNH archives (cf. Hol-
thuis & Sakai, 1970, p. 59) — said to be the only
species of monkey in Japan.
Inuus fuscatus: P. F. von Siebold, [1830-1842] (cf.
Temminck, 1842, p. 10; Holthuis & Sakai,
1970. p. 32; Aimi, 1992, p. 15)— unavailable
name used in two manuscripts in RMNH ar-
chives (Bemerkungen zu den Saugethieren von
Japan; Mammalium Japonicorum Enumeratio);
authorship of specific name attributed to [C. G.
C] Reinwardt (cf. Gijzen, 1938, p. 326).
Inuus fuscatus Gray, 1870, p. 32 — name pub-
lished as a junior synonym of Macacus specio-
sus "F. Cuv." (= I. Geoffroy, 1826) (for avail-
ability, see International Code of Zoological
Nomenclature, 1999, Arts. 11.6.1, 32.5.1);
"Mus. Leyderi" (= RMNH) cited as original
source of name; habitat, Japan. Blyth, 1875, p.
6 — conditionally proposed as technical name
for Japanese macaque, said to have been "for-
merly applied to it in the Leyden Museum."
Fooden, 1967, p. 251 — request for placement of
name on Official List of Specific Names in Zo-
ology. International Commission on Zoological
Nomenclature, 1970, p. 77 — name placed on
Official List of Specific Names in Zoology.
M[acacus] fuscatus: Sclater, 1 876, p. 332 — pre-
ferred specific name. Anderson, 1879, pp. 51,
78 — taxonomic history; external characters.
Thomas, 1906 ("1905"), p. 336 (not p. 361)—
three specimens reported; collector's field notes
published.
Mlacaca] fuscata: Pocock, 1926 ("1925"), pp.
1497, 1547 — spelling of generic name correct-
ed, gender ending of specific name changed ac-
cordingly. Fooden. 1966, p. 159 — proposed for
conservation as valid specific name.
[Macaca] fuscata-group: Miller, 1933, p. 5 — key
to external characters.
Macaca fuscata fuscata: Kuroda, 1940, p. 270 —
new rank; subspecies account.
Llyssodes] fuscatus: Pocock, 1921, pp. 225, 226
(part, reference to "Japanese macaque" only;
depicted specimen is misidentified Indochinese
stumptailed macaque) — new combination.
Lyssodes fuscata: Tate, 1947, p. 136 (part, distri-
bution also includes Yakushima) — revised gen-
der ending of specific name.
Lyssodes fuscata fuscata: Sowerby, 1943, p. 25 —
new combination. Furuya, 1962, p. 377 — der-
matoglyphics.
Pithecus fuscatus: Elliot, 1913, p. 195 (part, also
includes two Yakushima specimens) — new
combination; external characters.
[Silenus] fuscatus: Stiles and Nolan, 1929, p.
529 — new combination.
Inuus fucatus: [Temminck], [1827-1836] — un-
available name written in pencil on underside
of stand of RMNH 13-b (cf. Aimi, 1985, p. 24;
1992, p. 15), adult male, mounted skeleton (see
below. Remarks).
Innuus fucatus: H. Schlegel, [1830-1842] — un-
available name used in manuscript in RMNH
archives (Squelette de Japon).
Inuus specios[us] jap. Haberer, 1902, pp. 79, 82,
109 — ambiguously abbreviated subspecific
name, used in captions of skull illustrations.
Inuus speciosus japanensis Schweyer, 1909, pp.
1, 7, 9 — subspecific name proposed, possibly
inadvertently, in published doctoral disserta-
tion; based on 128 skulls obtained in Yokoha-
ma-city (118), Nikko (8), and Tokyo (2), pre-
served in collections in Munich, Berlin, and
Strasbourg.
[Macacus (Macacus)] yokoensis: Trouessart,
1904, p. 18 (part; not Matschie, 1900, p. 89)—
generic misidentification of African baboon;
specific name not available for Japanese ma-
caques.
Type Series — Fifteen or 16 specimens may be
included in the type series of Inuus fuscatus Gray,
1870 (p. 32). As explained below, these possible
syntypes include the following: one specimen for
which the name /. fuscatus was originally pro-
posed, but not published, by C. G. C. Reinwardt;
one or two additional specimens sent to RMNH
by Reinwardt; 12 specimens sent to RMNH by P.
F von Siebold and H. Burger; and one specimen
transferred from RMNH to BM(NH). According
to the International Code of Zoological Nomen-
clature (ICZN, 1999, Rec. 73F), a holotype should
not be assumed in cases such as this, where "it is
possible that the nominal species-group taxon was
based on more than one specimen."
For determination of the type series of /. fus-
catus Gray, three further provisions of ICZN
FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES
79
(1999) are relevant: (1) "The type series . . . con-
sists of all the specimens included by the author
in the new nominal taxon (whether directly or by
bibliogrophic reference)" (Art. 72.4.1. emphasis
added); (2) "any evidence, published or unpub-
lished, may be taken into account to determine
what specimens constitute the type series" (Art.
72.4.1.1); and (3) "[t]he type series of a . . . taxon
of which the name was first published as a junior
synonym . . . consists of the specimen (or speci-
mens) cited with that name in the published syn-
onymy" (Art. 72.4.3).
limits fuscatus was originally published as a ju-
nior synonym of Macacus speciosits. the Japanese
macaque, in Gray's (1870. p. 32) Catalogue of
Monkeys . . . in the Collection of the British Mu-
seum. This binomen subsequently was technically
made available — with Gray as author — when
Blyth (1875. p. 6) conditionally proposed it as a
replacement name for the taxon (see below. Re-
marks). The meager supplementary citation pro-
vided by Gray concerning the synonym /. fuscatus
is the appended abbreviation "Mus. Leyden."
which indicates that this name had been applied
to one or more Japanese macaque specimens at
RMNH (cf. Blyth. 1875. p. 6; Anderson. 1879. p.
78). According to ICZN (see above), the RMNH
specimen or specimens to which this name had
been applied constitute the type series of /. fus-
catus Gray. 1870.
An unpublished manuscript by R F. von Siebold
(see below. Remarks) establishes that the name /.
fuscatus was originally proposed by C. G. C.
Reinwardt to apply to a Japanese macaque that he
donated to RMNH; Reinwardt received specimens
from Japan in 1815-1822 during the course of his
government service in Java (Gijzen. 1938. pp. 46.
88). The specimen designated by Reinwardt as /.
fuscatus, which clearly is part of the type series
of /. fuscatus Gray, apparently no longer exists in
the collection of RMNH; it was not cataloged
there by either Schlegel (1876. p. 1 14) or Jentink
(1887, p. 27; 1892. p. 31).
In correspondence dated 20 January 1821.
Reinwardt indicated that he was shipping the skel-
eton of a Japanese monkey — "Simla {Cercoplthe-
cus) japonica" — to RMNH. and in correspon-
dence dated 12 May 1822. he indicated that he
was shipping another Japanese monkey skele-
ton— "Japansche aap" — to RMNH. Either — or
neither — of these specimens may be the same as
Reinwardt 's I. fuscatus specimen mentioned in the
preceding paragraph; neither apparently now ex-
ists in the RMNH collection. We are grateful to
C. Smeenk (e-mail message. 3 Dec. 2002), curator
of mammals at RMNH. for the cited information
concerning Reinwardt's 1821 and 1 822 correspon-
dence.
Judging from a penciled notation on the under-
side of the stand of RMNH skeleton no. 13-b (see
below. Remarks), the type series of /. fuscatus
Gray may also include some or all of 12 Japanese
macaque specimens that were present in the col-
lection of RMNH in or before 1867, when the text
of Gray's (1870. pp. [iii]. 32) work was printed
(cf. Schlegel. 1876. p. 114; Jentink. 1887. p. 27;
1892. p. 31). Of these specimens. 11 were col-
lected in 1823-1829 at unspecified localities by R
F. von Siebold. and one was collected in 1830-
1832 at an unspecified locality by H. Burger (cf.
Gijzen. 1938. pp. 297. 305; Holthuis & Sakai.
1970, pp. 38. 58). The 1 1 specimens collected by
von Siebold are no. 1-a. adult male, mounted skin
and skull (no. 7-d); no. 2-b. adult male, mounted
skin only; nos. 3-c and 4-d. adult females, mount-
ed skins only; no. 5-e. juvenile female, mounted
skin and mounted skeleton (no. 14-c); no. 6-f, ju-
venile female, mounted skin only; no. 8-e, juve-
nile male, skull only; nos. 9-f and 10-g, adult fe-
males, skulls only; no. 11-h, unsexed juvenile,
skull only; and no. 13-b, adult male, mounted
skeleton only. The specimen collected by Burger
is no. 12-a, adult male, mounted skeleton only.
Another possible member of the type series of
/. fuscatus Gray is BM(NH) 1842.1.19.95, adult
female (judging from long nipples), skin and
skull, locality unspecified (cf. Napier, 1981, p.
27); this specimen originally was included in the
RMNH collection. The BM(NH) register infor-
mation pertaining to this specimen is "Purchased
from J. Leadbeater [a dealer] who received [it] in
exchange from the Leyden Museum[']s [director]
M. Temminck." The specimen was received at
BM(NH) ca. 1842, and it is cataloged in Gray's
(1843, pp. X, 8) List of the Specimens of Mam-
malia in . . . the British Museum. The cited
BM(NH) register information was kindly provid-
ed to us by R Jenkins (e-mail message, 2 Jul.
2003), collection manager, mammals.
Although 14-15 specimens have been identi-
fied as possible syntypes of /. fuscatus (= M. f.
fuscata) Gray, 1870, we believe that, at present,
designation of a lectotype for this taxon would
serve no useful taxonomic purpose (cf. ICZN,
1999, Art. 74.7.3).
Type Locality — Japan; no further information
is available concerning collecting localities of the
syntypes.
80
FIELDIANA: ZOOLOGY
Distribution (Fig. 2B; Appendix 3) — Japanese
archipelago, from STN to 41.5°N, including the
large islands Kyushu, Shikoku, and Honshu and
the small islands Kojima, Kashima, Awajishima,
Shodoshima, and Kinkazan.
Diagnosis — Pelage on dorsal surface of trunk
pale yellowish brown to grayish brown to dark
golden brown; pelage on dorsal surface of hands,
pale yellowish brown to dark brown, not darker
than pelage on dorsal surface of trunk (Table 2).
Mean head and body length 522.8 mm in 53 adult
females, 570.1 mm in 41 adult males (Table 5);
mean sitting height 544.4 mm in 134 adult fe-
males, 596.9 mm in 99 adult males (Table 6).
Mean tail length 82.1 mm in 621 adult females,
89.8 mm in 277 adult males (Table 8). Mean
greatest length of skull 120.7 ± 4.56 mm in 184
adult females (extremes, 109.1-133.7 mm), 135.5
± 5.40 mm in 108 adult males (extremes, 121.3-
148.3 mm) (Table 19). Mean orbital height/
breadth index 1.00 ± 0.055 in 108 adult females
(extremes, 0.89-1.14), 0.95 ± 0.051 in 90 adult
males (extremes, 0.80-1.06) (Table 21).
Remarks — The oldest published species-group
name applicable to Japanese macaques, and hence
to the nominotypical subspecies, is Macacus spe-
ciosus I. Geoffroy, 1826 (p. 589) (Fooden, 1966,
p. 154). However, this name became widely mis-
applied to the Indochinese bear macaque; as a re-
sult, Macacus speciosus has been suppressed by
the International Commission on Zoological No-
menclature (1970, p. 77) and has been supplanted
by Inuus fuscatus Gray, 1870 (p. 32) (author and
date formerly cited as Blyth, 1875; see below).
The history of the use of fuscatus (Latin:
dusky) as a technical name for Japanese macaques
is somewhat complicated, and certain details re-
main obscure. According to a manuscript com-
posed ca. 1830-1842 by P. F von Siebold (see
following paragraph), the name Inuus fuscatus
was first used by C. G. C. Reinwardt to designate
a Japanese macaque specimen that Reinwardt do-
nated to RMNH. Although the date of this dona-
tion is unspecified, Reinwardt is known to have
shipped Japanese macaque specimens to RMNH
from Java during the period 1815-1822 (see
above. Type Series), when he served in Java as
director of agriculture, art, and science; prior to
serving in Java, Reinwardt had been the director
of the Government Cabinet of Natural History in
Amsterdam, and in 1823, after returning to the
Netherlands, he was appointed to a professorship
at Leiden University (Gijzen, 1938, pp. 5, 46, 88,
89, 326). We have been unable to locate Rein-
wardt's /. fuscatus specimen at RMNH, and no
such specimen is listed in the catalogs of Schlegel
(1876, p. 114) and Jentink (1887, p. 27; 1892, p.
31).
Von Siebold collected macaque specimens for
RMNH while serving as a physician in Japan dur-
ing the period 1823-1829 (Schlegel, 1876, p. 1 14;
Jentink, 1887, p. 27; 1892, p. 31; Holthuis & Sa-
kai, 1970, p. 25). After his service in Japan, von
Siebold took up residence in Leiden (1830-ca.
1846), where he composed the unpublished man-
uscript mentioned above. This manuscript, which
is titled "Bemerkungen zu den Saugethieren von
Japan," is preserved in the archives of RMNH
(copy of manuscript generously made available by
C. Smeenk, curator of mammals at RMNH; cf.
Aimi, 1992, p. 15). Although the text of the man-
uscript was written by von Siebold's secretary J.
J. Hoffman, von Siebold was responsible for the
contents. The manuscript presumably was written
between 1830, when von Siebold returned from
Japan to the Netherlands (Holthuis & Sakai, 1970,
p. 32), and 1842, when Temminck (1842, p. 10)
published a close French paraphrase of much of
von Siebold's German text. (Example: von Sie-
bold— "In der Menagerie des Sjogun zu Jedo be-
fand sich im Jahre 1 826 ein Albinos dieser Affen-
art, der in den Gebirgen von Nikko gefangen wor-
den war." Temminck — "On voyait en 1826, lors
du sejour de M. de Siebold au Japon, dans la me-
nagerie du Sjogun [I'Empereur] un albinos du
Speciosus, pris dans les forets en montagnes de
Nikko.")
Concerning authorship of the specific n?Ln\t fus-
catus, the section on Japanese monkeys in von
Siebold's manuscript contains the following sen-
tence: "Fine hochst merkwiirdige Abart dieses
Affen verdankt das Museum Herrn P. Reinwardt,
der dieselbe aus Japan erhalten und mit dem Na-
men I[nu]us fuscatus bezeichnet hat." (Transla-
tion: For a highly remarkable variety of this mon-
key, the museum is grateful to Herr P[rofessor?]
Reinwardt, who received it from Japan and des-
ignated it by the name Inuus fuscatus.) Von Sie-
bold also credited Reinwardt with authorship of
the specific ndimc fuscatus in another unpublished
manuscript — "Mammalium Japonicorum Enu-
meratio" — that is preserved in the RMNH ar-
chives; this manuscript includes the entry "Inuus
fuscatus Reinw. [= Reinwardt]."
A variant spelling of fuscatus appears in an un-
published list of von Siebold's specimens (RMNH
archives) that was prepared by H. Schlegel, then
a curator at RMNH (Gijzen, 1938, p. 53; Holthuis
FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES
81
& Sakai. 1970. p. 70): this list, titled '-Squelettes
de Japon," includes the entry Innuus fucatus"
(not "fuscatus"*). The same variant spelling. 'Tn-
uus fucatus."* also appears in a notation written in
pencil on the underside of the stand of one of the
monkey specimens collected by von Siebold —
RMNH 13-b. mounted skeleton of adult male (cf.
Aimi. 1985. p. 24; 1992. p. 15): this notation was
written by Temminck (C. Smeenk. RMNH. e-mail
message. 21 Jul. 2004). presumably sometime be-
tween 1827. when the first of von Siebold's spec-
imens arrived at RMNH (Holthuis & Sakai. 1970.
p. 54), and 1836. when Temminck (1836. p. xxii)
is known to have adopted speciosiis as the specific
name applicable to Japanese macaques (see
above. Synonymy).
In or before 1842, BM(NH) purchased a Japa-
nese macaque specimen (no. 1842.1.19.95) from
a British dealer who had acquired the specimen
from RMNH (see above. Type Series): this spec-
imen subsequently was cataloged as Macaciis spe-
ciosus by J. E. Gray (1843. pp. x. 8). then curator
of mammals at BM(NH). In or before 1 867. Gray
(1870. pp. [iii]. 31) again cataloged this specimen
as Macacus speciosus, but this time he indicated
that "Inuus fuscatus. Mus. Leyden" was a junior
synonym. His source for the supplementary indi-
cation is unknown: possible sources are the dealer
from whom BM(NH) 1842.1.19.95 was purchased
and/or RMNH staff members (cf. Gijzen. 1938. p.
311). Subsequently. Blyth (1875. p. 6) and An-
derson ( 1 879. p. 50) mistakenly asserted that Ma-
cacus speciosus was not applicable to the Japa-
nese macaque and adopted Inuus fuscatus or Ma-
cacus fuscatus as the name of this species (Food-
en, 1966. p. 156).
According to ICZN (1999. Arts. 11.6.1. 50.7).
Blyth's action made Inuus fuscatus available as a
specific name, with Gray as author and 1870 as
date of publication. Although Blyth's proposal
was conditional, this is irrelevant for works pub-
lished before 1966 (ICZN, 1999, Arts. 11.5.1.
15.1).
Specimens Examined — Total 1082: skins and
skulls. 25: skins only. 22: skulls or skeletons only.
1035 (Appendix 1).
Mticaca fusccUa yakui KuToda, 1940
Synonymy
Macacus fuscatus: Tliomas. 1906 ("1905"). p.
361 (not p. 336) — five specimens (two females.
three males) acquired by BM(NH) in 1905 from
A. Owston. collected in 1904; external charac-
ters, "dark in colour, but not darker than some
of the Shikoku examples."
Macaca fuscata: Kuroda. 1938. p. 112 (part) —
one Yakushima specimen listed; type history.
Lyssodes fuscatus: Kishida, 1953. p. 79 (part) —
taxonomic status, "the majority of Japanese
mammalogists in active service today do not
accept \\akui as a distinct insular subspecies]."
Macaca fuscata yakui Kuroda. 1940, p. 273, plate
47 — proposed as new subspecies; distribution
restricted to Yakushima: external characters.
Imaizumi. 1962. p. 10 — type history. Hill,
1974. p. 614 — type information. Napier. 1981,
p. 27 — five paratypes cataloged; taxonomic sta-
tus, "doubtfully distinct." Nozawa et al. 1991,
p. 431 — subspecific recognition questionable.
Kaneko and Maeda. 2002. p. 10 — type history.
Brandon-Jones et al.. 2004. pp. 99, 1 \4 — "rec-
ognition is doubtful."
Lyssodes fuscata yakui: Sowerby, 1943, p. 26 —
new combination. Furuya, 1962, p. 377 — der-
matoglyphics.
T^PE Series — The holotype of M. f yakui, by
original designation, is a juvenile male, skin
(?only). specimen no. 497. formerly in Nagamichi
Kuroda"s private collection in Tokyo. TTiis speci-
men was collected on Yakushima, 2 July 1905, by
A. Owston's collector (Nagamichi Kuroda, 1938,
p. 112: 1940. p. 273: Imaizumi. 1962. p. 10; Hill,
1974. p. 614). The specimen was destroyed by fire
during World War II in May 1945 (Austin et al.,
1948. p. 4: Imaizumi. 1962, p. 10: Nagahisa Ku-
roda. 1980. p. 24: Kaneko & Maeda, 2002, p. 10).
Although a small, separately housed part of Na-
gamichi Kuroda's collection escaped destruction
and subsequently was transferred to the Yamashi-
na Institute for Ornithology. Abiko. Chiba Prefec-
ture, we can confirm that the holotype of M. f.
yakui is not present in the mammal collection of
the latter institution (personal observation, assist-
ed by Collection Manager T Hiraoka, 20 Apr.
1999).
Six other specimens (paratypes) were included
by Kuroda (1940. p. 273) in his new subspecies
(International Code of Zoological Nomenclature,
1999. Art. 72.4). Of these, one was a living adult
(?female) (plate 47, animal, in color), then captive
at the Ueno Zoo, Tokyo: no part of this captive is
known to have been preserved. TTie remaining
five paratypes. cited by Kuroda from Thomas
(1906 ["1905"], p. 361). are skins and skulls in
82
FIELDIANA: ZOOLOGY
BM(NH) that were collected in 1904 by A. Ows-
ton as follows: nos. 1905.11.3.1 and 1905.11.3.2,
adult males, collected 4 October; no. 1905.1 1.3.3,
adult male, 18 October; no. 1905.1 1.3.4, adult fe-
male, 17 October; no. 1905.11.3.5, adult female,
18 October (cf. Napier, 1981, p. 27).
Type Locality — Yakushima (island), Kagoshi-
ma Prefecture, Japan. The exact place of collec-
tion of the holotype is unknown.
Distribution (Fig. 2b; Appendix 3) — Yakushi-
ma (island), Kagoshima Prefecture, Japan.
Diagnosis — Pelage on dorsal surface of trunk
dark golden brown; pelage on dorsal surface of
hands blackish, darker than pelage on dorsal sur-
face of trunk (Table 2). Head and body length
measurements not available; mean sitting height
510.1 mm in 34 adult females, 568.1 mm in 27
adult males (Table 6). Mean tail length 90.0 mm
in 44 adult females, 101.4 mm in 35 adult males
(Table 8). Mean greatest length of skull 113.4 ±
2.44 mm in 20 adult females (extremes, 109.6-
118.5 mm), 128.9 ± 5.23 mm in 29 aduk males
(extremes, 1 16.2-137.4 mm) (Table 19). Mean or-
bital height/breadth index 1.08 ± 0.065 in 20
adult females (extremes, 0.94-1.18), 1.06 ± 0.066
in 25 adult males (extremes, 0.96-1.20) (Table
21).
Specimens Examined — Total 182: skins and
skulls, 5; skulls or skeletons only, 177 (Appendix
1).
Evolution and Dispersal
Sixty-five species of nonvolant terrestrial mam-
mals are native to the island chain that extends
from Yakushima to Sakhalin (Table 53; Fig. 25).
Of these, 25 species (38%) are restricted in this
area to islands north of the Tsugaru Strait (be-
tween Honshu and Hokkaido; sill depth ca. 130
m), 27 species (42%) — including M. fuscata — are
restricted to islands south of this strait, and only
13 species (20%) occur both north and south of
the strait. The zoogeographic importance of the
Tsugaru Strait has been formalized by the desig-
nation of Blakiston's Line — which runs between
Honshu and Hokkaido — as a major faunal bound-
ary (Blakiston & Pryer, 1880, p. 177; Kuroda,
1939, p. 40; Tokuda, 1941, p. 133; Smith, 1983,
p. 463; Dobson, 1994, p. 99).
During the Pleistocene, the dispersal of non-
volant terrestrial mammals from the Asian main-
land to the Japanese archipelago presumably was
Table 53. Distribution relative to Tsugaru Strait
(between Honshu and Hokkaido) of Yakushima-Sakha-
lin species of nonvolant terrestrial mammals.'
Number Number
of
of non-
endemic endemic
Species
Yaku-
Yaku-
distribution
shima-
shima-
relative to
Salihalin Sakhalin
Tsugaru Strait
species
species
Totals
North only
1
24
25 (38%)
North and south
3
10
13(20%)
South only
19
8
27 (42%)
Totals
23
42
65(100%)
Reference: Dobson (1994, p. 96).
strongly controlled by the periodic appearance of
land bridges that were a consequence of cyclical
glacial advances and sea-level depressions. A re-
cent study indicates that the greatest sea-level
lowering during the last half-million years was ca.
- 140 m at 0.44 Ma (Rohling et al., 1998, p. 165).
The progenitors of mammalian species restrict-
ed to islands north of Blakiston's Line probably
were Palearctic species that dispersed during pe-
riods of low sea level from the Siberian mainland
to Sakhalin across the then-drained Tatar Strait
(sill depth ca. 15 m) (Fig. 25; Dobson, 1994, p.
99). Conversely, the progenitors of M. fuscata and
other species restricted to islands south of Blak-
iston's Line probably were Indo-Malayan species
that dispersed during periods of low sea level
from the Korean Peninsula area to Kyushu-West
Honshu across the then-drained Korea Strait (sill
depth ca. 130 m). Judging from morphological,
zoogeographical, and molecular biological evi-
dence, the mainland progenitor of M. fuscata
probably was similar or identical to M. mulatta
(Tables 25, 26; Nozawa et al., 1996, p. 24; Food-
en, 2000, p. 88).
The oldest macaque fossil unearthed in Japan
is an isolated lower third molar that is similar to
corresponding teeth in extant M. fuscata and is
tentatively dated at 0.43-0.63 Ma (see above. Fos-
sils). Korean macaque fossils, also similar to ex-
tant M. fuscata, are approximately the same age.
The successive development during this period of
two land bridges — now ca. 130 m underwater —
between the Asian mainland and southern Japan
is indicated by Japanese proboscidean fossils. A
working hypothesis — similar to that proposed by
Kamei (1969, p. 10) and Kamei et al. (1988, p.
FOODEN AND AIML SYSTEMATIC REVIEW OF JAPANESE MACAQUES
83
-j^p — I — I — I — I — I — \ — r
15 m*
Sakhalin
??
125^E
ISOT'E
135"E
Longitude
140'E
145'E
Fig. 25. Bathymetric map of Yakushima-Sakhalin island-chain area, illustrating probable land bridges exposed by
sea-level depression of 140 m; asterisks (*) indicate approximate sill depths of three zoogeographically important
straits (references: Keigwin & Gorbarenko, 1992. p. 347; Dobson. 1994, p. 99; Tada, 1994, p. 488; WORLDBATH
topography, 2000).
195; cf. Melnick et al.. 1993, p. 290; Nozawa et
al., 1996, p. 24) — is that M. fuscata is derived
from a M. mulatta-like ancestral population that
dispersed, during one or two glacial intervals ca.
0.43-0.63 Ma, from the Korean Peninsula area to
Kyushu-West Honshu via a now-submerged land
bridge (cf. Dobson & Kawamura, 1998, p. 390).
The progenitors of M. cyclopis may have dis-
84
FIELDIANA: ZOOLOGY
persed from the Asian mainland to Taiwan during
approximately the same geological period (Food-
en & Wu, 2001, p. 37).
Based on Y-chromosome DNA and mtDNA
variation in various macaque populations, Tosi et
al. (2003, p. 1431) have inferred that macaques
may have dispersed to Japan and Taiwan ca. 1.2
Ma. However, there is no fossil evidence for the
existence of macaques in either Japan or Taiwan
at this early date (cf. Dobson & Kawamura, 1 998,
p. 390), and the relevant oxygen isotope record
suggests that glacial sea-level depressions prior to
ca. 0.9 Ma would not have been adequate to cre-
ate requisite dry-land connections between the
Asian mainland and these islands (Shackleton,
1995, p. 245).
Macaca fuscata apparently has not been able to
disperse from Honshu to Hokkaido. This may be
because no land bridge has existed between these
two islands or — if a land bridge existed during an
exceptionally intense glacial interval — because of
cold climate prevailing in the region of the bridge
(Dobson, 1994, p. 100). During the last glacial
maximum (ca. 18 Ka), for example, in most of
northern Honshu (>35°N), broadleaf forest habi-
tats— suitable for M. fuscata — apparently were
less abundant than at present (Tsukada, 1982, p.
1092; Takahara et al., 2000, p. 673; Harrison et
al., 2001, p. 130).
During glacial sea-level lowering, M. fuscata
presumably was able to disperse to most or all of
the small shallow-water islands that surround
Kyushu, Shikoku, and Honshu. Subsequently,
however, this species apparently became extinct
on all except six of these small islands (Awajish-
ima, Kashima, Kinkazan, Kojima, Shodoshima,
and Yakushima). Such extinctions on small is-
lands have recently been recorded on Omishima,
Hiradoshima, and Tanegashima (see above. Geo-
graphic Distribution).
Subsequent to the initial arrival of M. fuscata
on the Japanese archipelago, its body size appar-
ently has tended to increase as the species dis-
persed northward, in accord with Bergmann's rule
(Figs. 5, 6, 9, 10, 15; Mayr, 1963, p. 320). Four
measurements of body size — sitting height, ante-
rior trunk length, hind foot length, and greatest
length of skull — generally tend to increase with
latitude from Yakushima to southern Honshu (ca.
35°N) but do not tend to increase with latitude
farther north on Honshu; one measurement of
body size — body weight — tends to increase with
latitude from Yakushima to northern Honshu. A
hypothesis to account for present body-size vari-
ation in M. fuscata is suggested by Kawamoto's
(2002, p. 68; see above. Molecular Biology and
Genetics) interpretation of mtDNA variation in
this species: body-size variation in M. fuscata
may have been completely Bergmannian prior to
the last glacial maximum, but northern popula-
tions (large-bodied) may have become locally ex-
tinct because they were unable to survive unfa-
vorable glacial climate conditions; during the sub-
sequent postglacial period, northern Honshu may
have been reoccupied by immigrants from south-
ern Honshu (smaller-bodied). Body weight may
be more plastic than the other four body-size at-
tributes, and it therefore may have more rapidly
regained a Bergmannian pattern of latitudinal var-
iation. The apparent reduction of body size in
Boso Peninsula and Kinkazan populations of M.
fuscata is unexplained.
In accord with Allen's rule (Mayr, 1963, p.
323), tail length in M. fuscata apparently has de-
creased relative to that in its presumed M. mulat-
r«-like progenitors (Tables 5, 8; Fooden, 2000, p.
29). Tail length in M. fuscata is approximately as
predicted by the tail length vs. latitude regression
determined by other fascicularis-group species
(Fooden & Albrecht, 1999, p. 433).
Pelage color in available samples of M. fuscata
generally becomes paler with increasing latitude,
in accord with Gloger's rule (Table 2; Mayr, 1963,
p. 324). This apparently is another example of ad-
aptation to local climate that has occurred since
monkeys initially arrived on the Japanese archi-
pelago.
As indicated above (see Molecular Biology and
Genetics), the history of dispersal of M. fuscata
within the Japanese archipelago that is implied by
mtDNA evidence differs from the history of dis-
persal that is implied by blood-protein evidence.
MtDNA evidence suggests that, since the last gla-
cial maximum, a relatively new M. fuscata pop-
ulation— with group II haplotypes — has rapidly
expanded its distribution in northern Honshu and
replaced a now disjunctly distributed older pop-
ulation— with group I haplotypes (Kawamoto,
2002, p. 68). Contrastingly, blood-protein evi-
dence suggests that extant monkey populations on
Yakushima, Boso Peninsula, and Shimokita Pen-
insula are relicts of a formerly widespread first
wave of immigrants to the Japanese archipelago
that subsequently were displaced elsewhere on the
archipelago by a possibly better adapted second
wave of immigrants (Nozawa et al., 1991, p. 433;
1996, p. 33). Further research is required to rec-
oncile these two interpretations.
FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES
85
In summary, the progenitor of M. fuscata prob-
ably was a M. mulatta-Uke population that dis-
persed via a now-submerged land bridge from the
Korean Peninsula area to Kyushu-West Honshu;
this dispersal probably occurred during one or two
glacial intervals of sea-level depression ca. 0.43-
0.63 Ma. In addition to dispersing to Kyushu,
Honshu, and Shikoku, M. fuscata probably also
dispersed to most or all of the surrounding small
shallow-water islands, but not to Hokkaido; sub-
sequently, the species apparently became extinct
on all but six of the surrounding shallow-water
islands. Since the arrival of M. fuscata on the Jap-
anese archipelago, its body size, tail length, and
pelage color apparently have tended to become
adapted to local climate conditions. As a result of
unsuitable climate and vegetation conditions dur-
ing the last glacial maximum (ca. 1 8 Ka), M. fus-
cata may have become locally extinct in northern
Honshu; during the subsequent postglacial period,
this area may have been reoccupied by a popu-
lation from southern Honshu.
Acknowledgments
For access to specimens and generous assis-
tance, we are deeply grateful to officials and staff
members of institutions cited above (see Intro-
duction). For valuable consultation, we thank Eric
Delson (AMNH), Siho Fujita (PRIKU), Goro
Hanya (PRIKU), Takashi Hiraoka (Yamashina In-
stitute for Ornithology), Mitsuo Iwamoto (JMC),
Paula Jenkins (BM[NH]), Yoshi Kawamoto (PRI-
KU), Naoki Koyama (Kyoto University), Shiro
Matsuoka (Working Group for Monkeys on Shi-
mokita Peninsula), Yukihisa Mito (JMC), Akio
Mori (PRIKU), Yasuyuki Muroyama (PRIKU),
Toyohiro Nishimoto (National Museum of Japa-
nese History), Hideyuki Ohsawa (PRIKU), Chris
Smeenk (RMNH), Shigeru Suzuki (Kyoto Uni-
versity), Kazuo Wada (Inuyama), Kunio Watana-
be (PRIKU), and Takao Yamaguchi (Kumamoto
University). Officials at Kozanji Temple gener-
ously permitted us to reproduce part of "Choju-
jinbutsu-giga" (Frolicking Animals and Figures
Scrolls). Photographs reproduced in Figures 3 and
1 1-14 were taken by Minoru Kinoshita (PRIKU).
W Burger, S. Drasner, M. Pannell, C. Simpson,
and H. Voris (all at Field Museum of Natural His-
tory) facilitated publication, and K. H. Hamilton
(Chicago) kindly assisted with translation of Jap-
anese literature.
The research of MA was supported by the Min-
istry of Education, Culture, Sports, Science and
Technology, Japan (10CE2005), and MEXT
Grant-in-Aid for the 21st Century COE Program
(A2 to Kyoto University). The research of JF was
supported by the Barbara E. Brown Fund for
Mammal Research.
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Appendix 1: Specimens Examined
(Total 1264)
Macaca fuscata fuscata, Total 1082
Skins and Skulls, 25
HONSHU, 3. Kanagawa (1): Fuji-san—
MNHN 1887/16. Mie (1): locality unknown—
MCZ 37709. Prefecture unknown (1): Kobe-
city— USNM 174730.
SHIKOKU, 6. Kochi (1): Chikaraishi— PRIKU
Coll. No. Shikoku 3. Tokushima (5): Jinrio —
BM(NH) 1906.1.4.1-1906.1.4.3; Kushigawa—
PRIKU Coll. No. Shikoku 1; Tachibana— PRIKU
Coll. No. Shikoku 2.
Island unknown, 16. AMNH 19137, 80038,
ANSP 1205, 6845, BM(NH) 1842.1.19.95,
1939.1050, MNHN 1886/101, 1929/548, RMNH
l-a/7-d/D20H, 5-e/14-c, 493, 1726, USNM
172412, 308426, 344996, ZMB 135/476.
Skins Only, 22
HONSHU, 9. Kanagawa (2): Fuji-san—
MNHN 1887/15, 1887/571 (skulls inside). Prefec-
ture unknown (7): USNM 13830 (female skin
mismatched with male skull; skin may belong
with skull USNM 13831), ZSBS 1901/101, 1904/
1718-1904/1722.
KOJIMA, 1. Miyazaki (1): KFL "Hiyoshi"
(name of monkey).
Island unknown, 12. ANSP 4952 (skull inside),
BM(NH) 1861.8.20.1, MCZ 6703, RMNH 2-b, 3-
C/D20G, 4-d/D20L, 6-f (skull inside), ZMB
12956/RexuCo, 22203, 32430, 34244, ZMUZ
11603.
Skulls or Skeletons Only, 1035
HONSHU, 863. Akita (1): Hachimori-machi—
PRIKU 6651 (mandible and postcranials only).
Aomori (4): Bushidomari — JMC 2577; Kusoudo-
mari — JMC 1757; Nishimeya-mura — JMC 1506;
Wakinosawa— PRIKU 2153. Chiba (284): Aika-
wa— PRIKU 6580, 6586, 6645, 6758, 6762; Fu-
kurogura,— PRIKU 6786; Futairi— PRIKU 6644
(cranial fragments); Hagiu — PRIKU 6588; Hina-
batake— PRIKU 6217, 6265, 6516, 6524, 6585,
6767; Hirooka— PRIKU 6283; Hongou— PRIKU
6761; Ishizuka— PRIKU 6493, 6507, 6522, 6774;
Kaisho— PRIKU 6258, 6287; Kamou— PRIKU
6242, 6280, 6282, 6290 (cranial fragments, man-
dible, postcranials), 6302, 6566, 6590; Kitako-
machi— PRIKU 6260, 6783; Kiwadahata— PRI-
KU 6293 (cranial fragments, mandible, postcran-
ials), 6300, 6495, 6593, 6765, 6773; Kokurano—
PRIKU 6256, 6276, 6284; Koshikiya— PRIKU
6776; Kozawamata— PRIKU 6294 (cranial frag-
ments, mandible, postcranials), 6624, 6756 (cra-
nial fragments), 6789; Kuradama— PRIKU 6235,
6236, 6241, 6245, 6274, 6291 (cranial fragments,
mandible, postcranials), 6292 (cranial fragments,
mandible, postcranials), 6296 (cranial fragments,
mandible, postcranials), 6298, 6299 (cranium re-
numbered "6239"), 6301, 6491, 6492, 6494,
6500, 6502, 6503, 651 1, 6519, 6573, 6587, 6591,
6592, 6594, 6607, 6608, 6752, 6757, 6768, 6769,
6778; Mera— PRIKU 6253; Kayano-Nanamaga-
ri— PRIKU 6251, 6501, 6625 (cranial fragments);
Nishihikasa— PRIKU 6497; Nutazawa— PRIKU
6760 (cranial fragments), 6779; Okugome — PRI-
KU 6505, 6510 (cranial fragments), 6576, 6599,
6600, 6643 (cranial fragments), 6754 (cranial
fragments), 6763, 6780, 6792; Okutani— PRIKU
6295 (cranial fragments, mandible, postcranials);
Okuyama— PRIKU 6513; Ooiwa— PRIKU 6496,
6499; Ootashiro— PRIKU 6243 (female, misla-
beled as male); Sanda— PRIKU 6575; Shigumi—
PRIKU 6244, 6271, 6787; Shimosakuma— PRI-
KU 6249, 6252, 6262, 6269, 6281, 6286, 6288,
6289, 6571, 6595, 6610; Shippara— PRIKU 6263,
6267, 6270, 6273, 6521, 6577, 6609, 6771, 6784;
Sunada — PRIKU 6237 (female, mislabeled as
male), 6266, 6272, 6297 (cranial fragments, man-
dible, postcranials), 6498, 6506, 6508, 6509,
6514, 6515, 6517, 6572, 6606, 6759, 6766, 6785,
6788, 6791, 6793; Tagura— PRIKU 6504, 6581,
6753, 6775, 6781; Takagoyama— JMC 2588
(mandible lacking), SU [1], [2], [3], [4], 5, 6, 7,
8, 9, 10, H-1, H-2, H-3, H-4 (postcranials only),
H-5, H-?, HK-34, I-l, 1-2, 1-3, 1-4, 1-5 (postcran-
FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES
105
ials only), I-6A, I-6B, I-6C, 1-9, I- 1 1, 1-12, 1-13
(postcranials only), 1-13, 1-14, I-? [1], I-? [2], K-
1 (postcranials only), K-2, K-3, K-4, K-5, K-11
or K-14 (postcranials only), K-12, K-13, K-15, K-
15?, Ko-1, Ko-2, Ko-3, Ko-4, Ko-5, Ko-6, Ko-7,
Ko-8, Ko-9, Ko-10, M-1, M-2A, M-2B, M-3, M-
4, M-5 (postcranials only), M-7, M-9, M-10, M-
11, M-1 2, M-1 3 (postcranials only), M-1 5, N-1,
N-2, N-3 (postcranials only), N-4, N-5, N-6, N-
6?, N-7 (postcranials only), N-8, N-9, N-10, N-
11, N-12B, N-12[A], N-13, N-14, N-16, N-17, W-
1, Y-1, Y-2, Y-2?, Y-3 (postcranials only), Y-4, Y-
5, Y-5?, Y-6 (postcranials only), Z-1, Z-2, "Jiro-
chou" (name of monkey); Tashiro — PRIKU 6257;
Tokuji— PRIKU 6770, 6790; Tsukagoshi—
PRIKU 6238, 6254 (cranial fragments, mandible
lacking, postcranials), 6255, 6259, 6279; Tsuki-
zaki— PRIKU 6247, 6250, 6264, 6268, 6518,
6523; Tsutsumori— PRIKU 6275, 6285; Utsutsu-
mi— PRIKU 6239, 6240, 6246, 6261, 6338, 6512,
6520, 6574, 6578, 6579, 6583, 6589, 6596-6598,
6755, 6764, 6777, 6782; Yamanaka— PRIKU
6582, 6772; Yokose— PRIKU 6248, 6277, 6278
(mandible fragmented); Yonezawa — PRIKU
6584. Fukui (15): Doumoto— PRIKU 5192, 5826
(cranial fragments); Hotokedani — PRIKU 5415
(postcranials only); Kaminaka-cho — PRIKU
5184, 5193, 5194; Kidani— PRIKU 5183, 5824
(cranial fragments, mandible, postcranials); Mi-
kata-cho— JMC 2012, 2034, 2035; Naka— PRI-
KU 5414, 5736 5737; Otomi— PRIKU 1869
(mandible lacking). Hiroshima (1): Miyajima-
cho — PRIKU 6647 (cranial fragments). Hyogo
(14): Kasuga-cho— JMC 3486, 3517, 3525, 3556,
3566, 3567, 3570; Sasayama-cho— JMC 3642,
3643, 3653, 3658, 3666, 3667; Takeno-cho— JMC
2502. Kyoto {\A): Hiyoshi-cho— JMC 3488, 3503,
3505, 3511, 3520-3522; Ine-cho— JMC 3501,
3504, 3515, 3519; Kamigamo— PRIKU 6129;
Mizuho-cho— JMC 3198; Wachi-cho— JMC
4214. Mie (4): Ilga-cho— PRIKU 257, 2111,
21 12; Oomiya-cho— PRIKU 6530. Nagano (261):
Aoki— PRIKU Coll. Nos. 86, 87; Atosugiyama—
PRIKU Coll. No. 49; Azumabasi— PRIKU 6317;
Buna — PRIKU 6106 (cranial fragments, postcran-
ials), 6189, 6200 (cranial fragments, mandible,
postcranials) , 6203, 6209; Fukagasawa— PRIKU
6477; Fukasawa— PRIKU Coll. Nos. 46, 47, 51;
Furukaidou— PRIKU 6488, 6568; Higashiina—
PRIKU 6461; Higashino— PRIKU 6098, 6113,
6474, 6480, 6800; Hinata— PRIKU 6099; Hina-
tayama— PRIKU 6163. 6817; Hodaka Golf
Course— PRIKU 6469; Horane— PRIKU 6463,
Coll. No. 178; Inokamireien— PRIKU 6820; Iri-
sawai — PRIKU 6799 (cranial fragments); Kaki-
zore— PRIKU 6470, 6619; Kamidaira— PRIKU
6100, 6105, 6115, 6167, 6168, 6177, 6330, Coll.
Nos. 65, 73, 78-81, 88-90, 108-115, 124, 141-
145; Kamimachi — PRIKU 6466; Kamiseinaiji —
PRIKU 6483, 6823 (cranial and mandibular frag-
ments); Karitayama — PRIKU 6490; Kashio —
PRIKU 6123, 6165; Kasugadaira— PRIKU 6124
(cranial fragments), 6180 (cranial fragments, post-
cranials), 6197 (cranial fragments, mandible, post-
cranials), 6208, 6214, 6487 (cranial and mandib-
ular fragments); Katsurashima — PRIKU 6806;
Kawanishi— PRIKU Coll. Nos. 55, 56; Kawasu-
so— PRIKU 6603; Kiso— PRIKU 1486, 2210; Ki-
tamura— PRIKU 6176/6454, 6192 (cranial frag-
ments, mandible, postcranials), 6194 (cranial frag-
ments, mandible, postcranials), 6199, 6201 (cra-
nial fragments, mandible, postcranials), 6202
(cranial fragments, mandible, postcranials), 6204,
6206, 621 1, 6454; Coll. No. 28; Konishi— PRIKU
6117; Kurouchi— PRIKU 6540; Kuwasononan-
bu— PRIKU Coll. Nos. 32-35; Maki— PRIKU
Coll. No. 118; Masugata— PRIKU 6464, Coll.
Nos. 173, 184; Metaka— PRIKU 6460; Minami-
machi— PRIKU Coll. No. 123; Misokawa— PRI-
KU 6564, 6565; Miyamoto— PRIKU Coll. Nos.
92, 97; Miyamura— PRIKU 6485, 6570, 6611,
Coll. Nos. 174, 185; Miyashiro— PRIKU 6121,
6179, 6339, 6458 (cranial and mandibular frag-
ments, postcranials), 6617 (cranial and mandibu-
lar fragments), 6673; Miyashita— PRIKU 6207;
Motomura— PRIKU 6459; Nakadachi— PRIKU
6615; Nakahora— PRIKU Coll. No. 94; Nakasu-
PRIKU 6322 (mandibular fragments, postcrani-
als); Nakayama— PRIKU 6796, Coll. No. 176;
Nezumiana— PRIKU 6104, 6344; Nishi— PRIKU
6801; Nishiharuchika— PRIKU 6122, 6174, 6182
(cranial fragments, postcranials), 6186 (cranial
fragments, mandible, postcranials), 6341 (cranial
fragments), 6455, 6468, 6484, 6671 (cranial frag-
ments), 6805, Coll. No. 134; Ochiai— PRIKU
6465, 6804; Ogikubo— PRIKU 6111, 6563, 6616
(cranial fragments); Oobara — PRIKU 6824; Ooji-
masan— PRIKU 61 16, 6210; Ookawara— PRIKU
6213; Oosaki— PRIKU Coll. Nos. 188, 189; Sa-
musawa — PRIKU 61 10, 6172 (cranial fragments),
6173, 6181 (cranial fragments, mandible, post-
cranials), 6187 (cranial fragments, mandible, post-
cranials), 6195 (cranial fragments, mandible, post-
cranials), 6196 (cranial fragments, mandible, post-
cranials), 6467, 6569 (note: external measure-
ments recorded for PRIKU 6187 and 6467 [Coll.
Nos. 98 and 99] appear to be mismatched with
the respective skeletons); Seto — PRIKU 6102,
106
FIELDIANA: ZOOLOGY
6212, 6456, 6618; Shiga-kogen— PRIKU 2141,
2142 (mandible lacking); Shimashima— PRIKU
6171, 6175, 6332, 6476, 6486; Shimizu— PRIKU
6166 (cranial and mandibular fragments), 6471,
6812 (cranial fragments), 6814; Shimoichiba —
PRIKU Coll. No. 71; Shimoseinaiji— PRIKU
6472, 6601, Coll. Nos. 175, 177; Shinyashiki—
PRIKU Coll. No. 23; Shiono— PRIKU 6614,
Coll. No. 182; Shouzawa— PRIKU 6479; Suza-
wa— PRIKU 6482 (cranial fragments); Takai—
PRIKU 6125, 6604, 6818, Coll. No. 180; Taki-
goshi— PRIKU 6479, 6922; Takinozawa— PRIKU
6794; Tatai— PRIKU 6531, 6648, 6649; Taza-
wa— PRIKU 6184, 6190, 6191 (cranial fragments,
mandible, postcranials). Coll. Nos. 190-193; Ten-
jinbara— PRIKU 6119, 6178, 6473, 6567, 6813;
Tokiwa— PRIKU 6331, 6612, 6613; Tsukiyo-
daira— PRIKU 6097, 6103, 6107, 6109, 6112,
6118, 6120, 6126, 6164, 6169, 6185, 6342, 6352,
6353; Ushimaki— PRIKU 6096, 6101, 6108,
6114, 6170, 6183 (cranial fragments, mandible,
postcranials), 6188 (cranial fragments, mandible,
postcranials), 6193, 6198, 6205, 6340, 6350,
6457, 6462, 6475, 6539, 6672 (cranial fragments),
6795, 6802, 6803, 6807-681 1, 6815, 6819, 6822;
Wakahowatauchi— PRIKU 6825; Wakamiya-
Hachimanjinja— PRIKU Coll. No. 130; Warabi-
taira— PRIKU 6489; Yakeyama— PRIKU 6797
(cranial fragments), 6798 (cranial fragments),
6816 (cranial fragments), 6821 (cranial frag-
ments), 6826 (cranial fragments); Yamabuki —
PRIKU 6481 (skull damaged); Yanagisawa— PRI-
KU Coll. No. 103; locality unknown, "Shinshiu"
(= Nagano Prefecture)— USNM 21116. Niigata
(5): Komogawa— PRIKU 6541, 6602, Coll. No.
Niigata 1 ; Takane— PRIKU Coll. Nos. Niigata 2-
3. Osaka (3): Minoo-city— PRIKU 277, 616, 617.
Shiga (1): Tsuchiyama-cho, Koka-gun [2] — TPM
(unnumbered). Shimane (156): Akashi — PRIKU
5721; Fukano— PRIKU 5149, 5179 (cranial frag-
ments, postcranials); Hasumi — PRIKU Coll. No.
Shimane 7; Higashihinobori— PRIKU 5154,
5176, 5178 (postcranials only); Hinagawa — PRI-
KU 6525, 6620, 6621, Coll. No. Shimane 10; Hi-
rasa— 561 1, 571 1 (postcranials only), 5713-5715;
Houin— PRIKU 5142 (cranial fragments), 5177,
5180; Imanishi— 5871, 5875, 5877, 5884 (cranial
fragments), 6131, 6138; Coll. No. Shimane 8;
Koujiro — PRIKU 5141 (cranial fragments, post-
cranials), 5143 and 5148 (postcranials only); Mik-
usu— PRIKU 5152, 5153, 5155, 5610: Mitoya-
cho— PRIKU 5612, 5710 (cranial fragments),
5739, 5867, 5880; Nishihinobori— PRIKU 5145,
5735; Okuyama— PRIKU 5741 (female, misla-
beled as male); Shimogou — PRIKU 5136 (po-
stcranials only), 5137, 5138, 5139 (postcranials
only), 5140, 5144, 5147, 5156, and 5238 (4 spec-
imens, postcranials only), 5239 (cranial frag-
ments, postcranials), 5240 (postcranials only),
5712, 5716, 5720, 5827 (postcranials only),
5859-5863, 5865, 5868-5870, 5878, 5883, 5885,
5886, 5889, 5890, 5892, 5893, 5895, 5980 (cra-
nial fragments), 6142, 6152, 6160, 6351; Taku-
wa— PRIKU 5151, 5708 (mandible broken),
5709, 5722, 5881 (cranial and mandible frag-
ments), 6132, 6143, 6529; Tsukinoya— PRIKU
5146, 5150 (mandible and postcranials only),
5241 (cranial fragments, postcranials); Ueda —
PRIKU 5864, 5872-5874, 5882, 5887, 5888,
5891, 6127, 6128, 6130, 6133-6137, 6139, 6146,
6148-6150, 6153-6155, 6158, 6161, 6438, 6526,
6527; Ushiroyama— 5738, 5876, 5894, 5979,
6140, 6141, 6144, 6145, 6147, 6151, 6156, 6157,
6159, 6162, 6528, Coll. Nos. Shimane 1-6, 9, 1 1-
24; Yoshida-mura— PRIKU 5866. Shizuoka (5):
Hagachizaki— PRIKU 1381; Hamakita-city—
PRIKU 1376, 1377, 1380; locality unknown—
PRIKU 6670. Tochigi (39): Imaichi-city— TPM
59/171, 61/074, 61/118-61/121, 61/123, 61/132;
Kuriyama-mura — TPM 61/090, unnumbered skull
(mandible lacking); Nikko-cityTPM 56/083-56/
086, 57/095, 57/178, 58/067, 58/071, 59/135, 59/
170, 60/101, 60/102 (mandible lacking), 60/106,
60/107, 60/246, 61/067, 61/072, 61/080, 61/091,
61/092, 61/1 10, ZMB Coll. Nos. III-X. Prefecture
unknown (56): "Jedo'V'Yedo" = Tokyo — ZMB
A3469, A3470, A3476, A3481, 5982, unnum-
bered female skeleton acquired 13 Feb. 1874;
"Jokohama" = Yokohama-city (cf. Schweyer,
1909, p. 8)— AIUM 3472-3480, 3482-3487,
3489, 3490, 3492-3506; "Tokyo Market"—
USNM 13830, 13831, 20937; "Yokohama"—
ZSBS 1902/97; locality unknown— ZSBS 1901/
69, 1901/71-1901/83.
KINKAZAN, 15. Miyagi (15): MUE K/1, K/2,
K/6, K/lO-K/15, K/20, K/24, K/28, K/33, K/Yl,
K/Y6.
KOJIMA, 47. Miyazaki (47): KFL 1, 2 (crani-
um broken), 3-9, 10 (mandible lacking), 11, 12,
13-15 (mandibles lacking), 16, 17 (mandible
lacking), 18, 19 (mandible lacking), 20, 21, 22
(mandible lacking), 23 (cranial fragments), 24, 25
(mandible lacking), 26, 28 (cranial fragments),
29-34, 35 (cranial fragments), 36-45, "Usagi"
(name of monkey), unnumbered skull (mandible
lacking), unnumbered mandible only.
KYUSHU, 18. Kagoshima (5): Kanayama—
PRIKU, five uncatalogued skulls. Miyazaki (1):
FOODEN AND AIMI: SYSTEMATIC REVIEW OF J7\PANESE MACAQUES
107
Toimisaki— JMC 1747. Oita (12): Takasakiya-
ma.— JMC 1949. 3971. PRIKU 2131. 2133. 2143.
2146. 2152. 2154 (mandible and postcranials
only). 2155-2158.
SHODOSHIMA. 47. Kagawa (47): JMC 925.
PRIKU 1405-1409. 1410 (mandible only). 1411
(mandible and ulna only). 1412 (mandible and
postcranial fragments only). 1413-1421. 1429
(mandible and postcranials only). 1431, 1432
(cranial fragments). 1433 [1]. 1433 [2]. 1434.
1438 [1]. 1438 [2]. 1438 [3]. 1444. 1446-1448.
1450 [1], 1450 [2], 1455, 1456 (mandible and po-
stcranials only). 1457 (mandible and postcranials
only). 1458-1460. 1464-1466. 1468. 1470. 1472-
1474.
Island unknown, 45. AIUZ 2063. 6064. AMNH
35640. BM(NH) 1850.8.15.2. NHR Z2266.
Z2267. PRIKU 4715. 4717. and 4718 (3 speci-
mens, postcranials only). 5879 (juvenile skull
mistakenly associated with data pertaining to a
large male). 6215. 6216. 6218. 6303-6305^ 6306
(cranium broken). 6307-6312. 6467 (adult female
mistakenly associated with data pertaining to an
infant male), unnumbered skull (with note refer-
ring to PRIKU 5139 and PRIKU 5140). RMNH
8-e. 9-f. 10-g. 11-h. 12-a. 13-b. USNM 259982.
260229, 344995. ZMB 12957. 25547. 25548.
25550. 25552-25554, RexuCo, unnumbered male
skeleton acquired in 1874. ZMUZ 1 1603a. ZSBS.
unnumbered male skull.
Macaca fuscata yakui, Total 182
Skins and Skulls, 5
YAKUSHIMA. 5. Kagoshima (5): locality un-
known—BM(NH) 1905.1 1.3.1-1905.1 1.3.5.
Skulls or Skeletons Only, 177
YAKUSHIMA, 177. Kagoshima (177): Hany-
ama— PRIKU 1571. 1572. 1729. 1730. 1731
(mandible lacking). 1732 (mandible lacking).
1733. 1749. 3430,^3957-3959, 4395 (cranial frag-
ment). 4490. 4491. 5424 (skull broken). 5425,
6626. 6627 (mandible and postcranials only),
6628-6630. 6631 (mandible lacking). 6632, 6633.
6634 (mandible and postcranials only). 6635,
6636 (mandible fragment only), 6637-6642.
6751; Hinokuchi— PRIKU 3153 (male skeleton
with mismatched female mandible). 3154-3157;
Issou— PRIKU 3158; Kamiyaku-cho (30°16'N)—
PRIKU 2148, 3433-3446; Kojima— PRIKU 2610
(cranial fragments, mandible, postcranials) ; Kun-
iwaridake — PRIKU 2167 (cranial fragments,
mandible, postcranials; Kusugawa — PRIKU
2595: Miyanoura— PRIKU 2147 (mandible lack-
ing); Mugio— PRIKU 2588. 2591. 2594. 2607,
2609. and 2611-2615 (7 specimens, postcranials
only); Nagata — PRIKU 2168 (mandible lacking).
2585-2587. 2608 (postcranials only). 3159-3162;
Nakama— PRIKU 2593. 2597-2605. 2616-2618
(3 specimens, postcranials only); Nakase — PRI-
KU 2606 (postcranials only); Nakasegawa — PRI-
KU 2584. 2589. 2590; Seibu-rindou— PRIKU
3351 (mandible lacking). 3352 (cranial and man-
dibular fragments): Shichigodake— PRIKU 2592.
2596; Shirakoyama— PRIKU 3141-3144. 3145
(postcranials only). 3146. 3147. 3148 [1] (infant
skeleton with mismatched juvenile mandible),
3148 [2] (aduh skeleton, skull damaged). 3149,
3150 [1], 3150 [2], 3150 [3]. 3151 (cranial frag-
ments). 3152 [1] (skull damaged). 3152 [2], 3152
[3]. 3152 [4], 3152 [5], 3451-3453; Yudomari—
PRIKU 3123-3131. 3132 (postcranials only),
3133-3140; locality unknown— JMC 601, PRIKU
3447-3450. 3454-3456. 3457 (cranial fragments),
4120 (cranial fragment and mandible only). 41 21-
4124, 4459 (cranial fragment), 4460 (skull dam-
aged), 4461, 4462, 4463. and 4720 (3 specimens,
postcranials only). 4941. 5009. 5227, 5247-5249,
5373 (postcranials only). 5416. 6083-6086. 6951,
6952, 6953 (cranial fragments, postcranials),
6954. 6955.
Appendix 2: Macaca fuscata
Localities Reported in Postal Survey
Conducted by K. Hasebe in 1923
(Fig. 2A; cf. Iwano, 1974, p. 5;
Uehara & Koganezawa, 1976, p. 742;
Amagasa & Ito, 1978, p.96); for
Locality Details, see Gazetteer,
Appendix 4.
YAKUSHIMA: Kagoshima Prefecture— Yakxi-
shimayama.
TANEGASHIMA: Kagoshima Prefecture— Fu-
rutayama et al.; Masudayama.
KOJIMA: Miyazald Prefecture — Kojima.
KYUSHU: Kagoshima Prefecture — Bono-mi-
saki; Shimoyama; Shibisan; Shiraki; Harimochi;
Kirishimasanrin; Onomi and Otaniyama; Takaku-
madake, Kanoye-cho; Kamimobiki vicinity; Taka-
kumadake; Takakuma; Kunimidaira vicinity; Ina-
108
FIELDIANA: ZOOLOGY
odake vicinity; Sarutsubo vicinity. Miyazaki Pre-
fecture— Misaki-Makinouchi vicinity; Ariki Na-
tional Forest vicinity; Kirishimayama,
Nishidake-son; Kirishimayama vicinity, Takahara-
son; Ohira National Forest vicinity; National For-
est, Suki-son; Ichifusa vicinity; Tatsufusayama vi-
cinity; Fukiyama National Forest; Kanasumi Na-
tional Forest, Kijo-son; Osuzu National Forest;
Tedayama vicinity; Nishinoyakaeyama; Totoro-
gauchi vicinity, Kadokawason; Okumiyama; To-
torogauchi, Kitagouson; Mukabakiyama; Hinoki-
yama; Ookueyama vicinity; Sokujitsunomine; Na-
misegawara; Tsuriganeyama; Higakureyama vi-
cinity; Oshioi; Yumigiyama. Oita Prefecture —
Mumeinoyama, Onoichi-mura; Mumeinoyama,
Shigeoka-mura; Odairayama vicinity; Katamuki-
yama; Mitakeyama; Honjoyama; Mumeinoyama,
Imaichi-mura; Mumeinoyama, Kawanobori-mura;
Taninoshiriyama; Inouesanrin; Onagarayama For-
est; Shihatsuyama; Shikakumadake, Shimogou-
mura; Shikakumadake, Yamautsurui-mura; Shigi-
ra Shinyabakei; Hikosanroku; Ota; Tano; Futago-
yama; Higashi- and Nishi-Kurokiyama. Kuma-
moto Prefecture — Nishiohira; Gongenyama
vicinity; Shiroiwayama; Yamate vicinity; Shira-
kiyama vicinity; Sakasegawa vicinity; Kosadake;
Fukabayama, Fukuoka Prefecture — Kusenbuya-
ma vicinity; Kawaradake, Kawara-machi; Egawa
vicinity.
KASHIMA: Ehime Prefecture — Kashima.
SHIKOKU: Ehime Prefecture — Aikawayama;
Shigekiyama; Masakiyama; Sasagou vicinity; Shi-
monaru; Kurase vicinity; Tataki vicinity; Higashi-
mikatagamori; Nakao; Kitamine; Amagoitaki wa-
terfall vicinity; Kuragariyama; Tatsukawayama;
Daieiyama; Tanegawayama vicinity; Fujigaishi vi-
cinity; Higashinokawa vicinity; Somakawa-mura;
Gayatani vicinity; Ibushikanyama vicinity; Koya-
yama; Ondayama vicinity; Akago; Umegatani vi-
cinity; Ryuoka-mura National Forest; Okuyama.
Kochi Prefecture — Kashiwajima; Souro vicinity;
Onozumi vicinity; Kurokawa; Kunimi vicinity;
Nakahama vicinity; Tsuro vicinity; Shimonokae
vicinity; Kujuu vicinity; Yamada; Kyouhou vicin-
ity; Fujikawa vicinity; Nagaoi; Tsuneroku vicini-
ty; Nakatsugawa vicinity; Nonogawa vicinity;
Kannonyama; Nadanishiyama vicinity; Nadayama
vicinity; Matsubakawayama vicinity; Amagoiya-
ma; Tachibanaseiryuu National Forest; Tsubaya-
ma; Yasui National Forest; Nakakurayama; Taka-
bagamori; Kirakuyama vicinity; Seto; Kiyano vi-
cinity; Minamikawa vicinity; Tochu vicinity; Shi-
mokawaba vicinity; Shiragayama; Ouyama
vicinity; Nishitoyonaga-mura; Makitaniyama vi-
cinity; Okugataniyama; Chaen; Nonesankei; Mo-
toyama; Oozakurayama; Yokootsuzukiyama vi-
cinity; Sugawayama vicinity, Iketaniyama vicini-
ty; Kanegaryumorisankei; Iwataniyama. Tokushi-
ma Prefecture — Kushigawa vicinity; Hirai
vicinity; Kitousanrin, Okukitou-son; Kitousanrin,
Kamikitou-son; Shindenyama; Kigiura vicinity;
Senbagatake vicinity; Akamatsu; Kawachi vicin-
ity; Nadayama; Gongendaki vicinity; Kakeban vi-
cinity; Fukuhara-son; Yanagitani; Okutachikawa
vicinity; Shikakubidani; Ooboke; Tokuzen-nishi
vicinity; Motoi vicinity; Kashiji vicinity; Kuchi-
yama-son; Iwaya; Handairayama-son; Yuumaya-
ma; Kawatayama vicinity; Koyadaira; Kidooku;
Nagafuka vicinity; Hanto-cho; Kitanada-son;
Gosho-son. Kagawa Prefecture — Yamawakiya-
ma; Higashiyama National Forest; Asan Mountain
Range.
SHODOSHIMA: Kagawa Prefecture— Tmma-
san; Kensozan, Oonude-mura; Kensozan, Kitaura-
mura; Kensozan, Oobe-mura; Kensozan, Kusaka-
be-mura; Kensozan, Yasuda-mura.
OMISHIMA: Yamaguchi Prefecture — Omishi-
ma, Senzaki-cho.
AWAJISHIMA: Hyogo Prefecture— Yuzuru-
hayama; Kashiwabarayama foothills.
HONSHU: Yamaguchi Prefecture — Onigajo;
Gezan; Sarugababa; Shirataki; Nagatokyo vicini-
ty, Kawakami-son; Nagatokyo vicinity, Ikumo-
son; Nagatokyo vicinity, Shinobu-son; Hagi-ma-
chi; Northern villages, Tsuno-gun; Kuratani vicin-
ity; Kuroda vicinity. Hiroshima Prefecture — Go-
kurakujisanrin; Nenbutsudani National Forest;
Onosanrin; Masakiyama; Yokogawaburaku vicin-
ity; Karioyama vicinity; Takiyamagawa vicinity;
Ryuuzuyama; Samuhikiyama; Kabetouge vicinity,
Honji-son; Kabetouge vicinity, Minamikata-son;
Doutokoyama; Hongushisan; Shirokiyama vicini-
ty; Takayama; National Forest, Nishishiwa-son;
Shirokiyama; Takanosuyama, Kuba-son; Takano-
suyama, Takeni-son; Takamurayama, Nyuno-son;
Youkurayama, Zen-nyuuji-son; Youkurayama,
Kamikitakata-son; Youkurayama, Funaki-son;
Youkurayama, Ookawa-son; Buttsujisan; Koni-
shioku; Ootaniyama vicinity; Ryuozan, Miyauchi;
Kugayama National Forest vicinity; Dakeyama;
Tarekaya Natioanl Forest; Iwamiyama vicinity;
Hizaoyama; Taishakugawa vicinity; Inyou Moun-
tain Range vicinity, Kimita-son; Inyou Mountain
Range vicinity, Funo-son; Inyou Mountain Range
vicinity, Sakugi-son. Okayama Prefecture — You-
zenotaki; Abeyama; Kanehirayama; Tenjinyama;
Sarugamiyama; Tsuchiosanrin vicinity; Furutani
National Forest; Douyama; Kamikumatani, Ku-
FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES
109
matani-son; Karamatsu; Akoumayama; Kanba;
Yumoto; Gagyusan, Tsugawa-son; Gagyusan,
Takahashi-cho; Gagyusan, Matsuyama-son; Ita-
mayama; Kuniyama National Forest; Takiyama;
Okutsugawa National Forest. Shimane Prefec-
ture— Kaneyamadani; Tachigouchi; Tadeno; Ka-
mitakashiri; Kabatani; Shiroyama, Hatagasako-
mura; Shiroyama, Tsuwano-cho; Tadachiokuya-
ma; Yokomichi vicinity; Takitani; Sumikawa vi-
cinity; Rakanyama, Hiromi; Kametaniyama
vicinity; Itotaniyama; Kuigautsu, Hinui-mura;
Maruseyama; Kirigataniyama vicinity; Iwaidani-
yama; Kanmuriyama; Kurokagara vicinity; Ooe
Takayama; Soratani; Hamahara-mura; Akutaya-
ma; Tsunotani; Kogarashigataniyama vicinity;
Ooyahongou; Yashiroyama, Yashiro-mura; Yata-
kiyama vicinity, Nishida, Yusato-mura; Ginzan
vicinity; Oshihara-mura; Hata-mura; Matsukasa-
mura; Yoshida-mura; Tai-mura. Tottori Prefec-
ture— Okinoyama; Momobayama; Iwayadou, Ike-
da-son. Hyogo Prefecture — Kusaki-mura; Miu-
chisanrin, Ishii-mura; Funakoshiyama; Kawaraya-
ma; Nabegatani; Akanishi vicinity; Rengeiwa
vicinity; Kayano vicinity; Kishida; Yukihikoya-
ma; Nagakura vicinity, Hase-mura; Kurataniyama,
Tataragi, Nakagawa-mura; Kanatokoyama, Itoi-
mura; Myomisan; Nishitokoo vicinity; Kanato-
koyama, Aihashi-mura; Mikuniyama; Houtaku-
san; Egasayama; Sasagamine; Atagoyama vicini-
ty; Akaioku, Nunuki-mura; Ishidosan; Mitsudake;
Ikuno, Doujou-mura; Kitayama, Shioze-mura;
Mayasan. Osaka Prefecture — Minoosan National
Forest; Hattasan vicinity; Katsuojisan; Shakaga-
dake; Ushitakisan; Inunakisan. Wakayama Prefec-
ture— Onoueyama; Ui, Ishigaki-mura; Otayama;
Nikkosan; Nakaodani vicinity, Go-mura; Muro-
kawayama; Kashiyama vicinity; Kawanaka-mura;
Hidaka Mountain Range; Kawamatasanrin; Shi-
mosanji-mura; Kamisanji-mura; Higashiyama vi-
cinity, Kamihaya-mura; Ryujinyama vicinity;
Takaosan vicinty, Nagano-mura; Toragamine
Mountain Range vicinity; Mochihirasan; Takao-
san vicinity, Chikano-mura; Ryujinyama vicinity,
Inari-mura; Ryujinyama vicinity, Kamiakitsu-
mura; Jujotouge vicinity; Mugikomoriyama vicin-
ity; Damasanchi, Ichinose-mura; Ashiyama vicin-
ity, Ikuma-mura; Shokawagoe vicinity; Hirutani
vicinity; Kongoumine vicinity; Endani vicinity;
Mimaisan vicinity; Futagoyama vicinity; Nakano-
goudani vicinity; Ebitadani vicinity; Shimizuyama
vicinity; Mureyama vicinity; Shigatakiyama vi-
cinity; Arida-mura/Higashimuro-gun boundary;
Fujihashi-mura/Higashimuro-gun boundary; Nyu-
douzan vicinity; Miokawa-mura; Shichikawa-
mura; Ogawa-mura; Takaike-mura; Hachirouya-
ma; Kamioota-mura; Mizunoshitayama; Irokawa-
mura; Takata-mura; Kitanokawa vicinity; Yo-
mura; Sunomata vicinity, Ukegawa-mura;
Karekidani, Shikiya-mura; Mikoshi vicinity, Mi-
sato-mura; Kujuu-mura; Tamakikuchi-mura. Nara
Prefecture — Kitamatasanrin, Nosegawa-mura;
Akataniyama; Hirosesanrin, Tenkawa-mura; To-
tsukawa-mura; Shimokitayama-mura; Kamikita-
yama-mura; Kawakami-mura; Mitsue-mura; Beni-
gadake vicinity; Taishiyama, Murou-mura; Taka-
toriyama National Forest, Takatori-cho; Takatori-
yama National Forest, Takaichi-mura. Kyoto
Prefecture — Konoshiro, Takano-mura; Oyama vi-
cinity; Choenzan; Taikoyama; Ookaneyama;
Houzawayama; Koryujiyama; Akaiwayama, Oka-
danaka-mura; Akaiwa Mountain Range, Maruyae-
mura; Kagamisakadani, Oono-mura; Sumugitani,
Miyajima-mura; Horatani vicinity, Tsurugaoka-
mura; Ashiu vicinity, Chii-mura; Hacchoyama;
Kumamidani vicinity, Kuroda-mura; Takitani vi-
cinity, Kuda-mura; Chishorodani; Urayama, Shi-
no-mura; Arashiyama vicinity; Yanagitani, Kaiin-
ji-mura; Makiosan vicinity; Ujitahara-mura. Mie
Prefecture — Tadosan vicinity, Komi-mura; Tado-
san vicinity, Tado-mura; Tashida vicinity. Hatta-
mura; Toriidoyama; Yunoyama; Kanmuriyama;
Nyudougatake; Sengatake; Nonoboriyama vicini-
ty; Nakatsugawayama; Kabutoyama; Shakujou-
gadake; Hiraki vicinity, Nagano-mura; Takarajou
vicinity, Awa-mura; Shorenjisan vicinity; Naga-
saka-mukaiyama; Iwayama, Tarou-mura; Ohara-
dake; Idodani vicinity; Hirakura, Yahata-mura;
Kuchisubosanrin; Awano, Kawamata-mura;
Hachisusanrin vicinity, Mori-mura; Miyama Na-
tional Forest, Ookawachi-mura; Shimatani vicin-
ity, Ryounai-mura; Aritani, Hagiwara-mura; Oo-
sugitani-mura; Oouchiyama-mura; Kasagi, Kashi-
wazaki-mura; Kurosu vicinity, Shimazu-mura;
Fujikiya forest, Nanaho-mura; Ishigagawa, Naka-
gawa-mura; Goriyama, Ogawagyou-mura; Higa-
shimiyamata vicinity, Ichinose-mura; Oshibuchi
vicinity, Hobara-mura; Michikata vicinity, Naka-
jima-mura; Oitsuge vicinity, Nishiki-mura; Nisa-
kasanrin vicinity; Tochiyama vicinity; Shimaka-
chisanrin vicinity; Ooike beach; Ookawachiyama
vicinity; Iwaidani vicinity; Wataka vicinity, Owa-
se-cho; Higashimisaki vicinity; Morimatsu vicin-
ity, Kitawauchi-mura; Sone vicinity, Minamiwau-
chi-mura; Komataoku vicinity, Asuka-mura; Mo-
mozaki vicinity, Isato-mura; Ogawaoku vicinity,
Kamikawa-mura; Komori, Nishiyama-mura;
Oomine vicinity; Wake vicinity, Kamikawa-mura;
Oosato, Onodani-mura; Asaoki, Mifune-mura;
110
FIELDIANA: ZOOLOGY
Matani, Agira-mura. Shiga Prefecture — Iwamaya-
ma; Hieizan; Hirasan vicinity, Kido-mura; Hira-
san vicinity, Katsuragawa-mura; Komatsu-mura;
Tsunogawa-Akana vicinity; Fukashimizu vicinity,
Kawakami-mura; Hidarimaeyama vicinity; Cho-
shigatani; Kataoka-mura; Niu-mura; Sugino-
mura; Ibukiyama, Ibuki-mura; Ibukiyama, Suijou-
mura; Ryozenyama, Kashiwabara-mura; Ibukiya-
ma, Samegai-mura; Ryozenyama, Toriimoto-
mura; Ryozen, Seritani-mura; Wakigahata-mura;
Takidani vicinity, Ootaki-mura; Higashiogura-
mura; Akasugiha vicinity, Yamakami-mura; Oo-
gawara, Aiga-mura; Kurotaki, Yamauchi-mura;
Tomikawayama. Fukui Prefecture — Aonogou-
mura; Takahama-cho; Saburi-mura; Wada-mura;
Hongou-mura; Ooshima-mura; Kado-mura; Hitot-
sudai, Okunata-mura; Nishiogawa vicinity, Uchi-
tomi-mura; Ayuhanasanrin, Nishida-mura; Mika-
tasanrin, Ya-mura; Shinjou vicinity, Mimi-mura;
Kurokawayama; Okuaso vicinity, Arachi-mura;
Takasuyama; Ashimi-mura; Ohara vicinity, Kita-
tani-mura; Gokairiaiyama, Kamishou-mura; Ni-
gure, Kamianama-mura. Ishikawa Prefecture —
Kuratani vicinity, Saigawa-mura; Okuike vicinity,
Kawachi-mura; Jadani vicinity, Yoshinotani-
mura; Hidaritsubute vicinity, Torigoe-mura; Ozoe-
chinai, Okuchi-mura; Kuwashimachinai, Shira-
mine-mura. Aichi Prefecture — Hongusan, Ichino-
miya-mura; Toyamagoryouchi vicinity, Nagashi-
no-mura; Ebi-cho; Kandayama; Myojinyama;
Ooiriyama, Sono-mura; Yatsudake vicinity; Ha-
guro-mura; Kurisuyama; Hongusan, Gakuden-
mura; Owarifuji vicinity. Gifu Prefecture — Tado
Mountain Range, Ishizu-mura; Tado Mountain
Range, Shiroyama-mura; Tokiyama; Oobora-Sa-
tani, Tara-mura; Shogatake vicinity; Ootani, Ma-
kita-mura; Shimonotani vicinity, Sakauchi-mura;
Shiratani vicinity; Kouchi, Yokokura-mura; Ooka-
wara vicinity, Neo-mura; Kinbaradani, Toyama-
mura; Kamagatani, Kamiijira-mura; Okuookura
vicinity, Kuzuhara-mura; Natsusaka vicinity, Ki-
tayama-mura; Tarudaira vicinity, Horado-mura;
Kawaurayama, Itadori-mura; Shimoyama vicinity,
Shimomaki-mura; Kouwayama vicinity. Suhara-
mura; Akakuzureyama, Shimonoho-mura; Otsu-
kamebora, Nakanoho-mura; Hatofukiyama; Fu-
kubegadake; Nahisanrin, Aioi-mura; Konahisan-
rin vicinity, Nishiwara-mura; Negitani, Nishika-
wa-mura; Inunaki, Kuchimyougata-mura;
Gozenyama; Sangenyama; Senchougahara; Hige-
tayama vicinity, Takane-mura; Byobuiwa, Kado-
hara; Kanakidoyama; Fukatani vicinity, Yama-
nokuchi-mura; Jigokudani vicinity, Siioukawa-
mura; Aratani vicinity, Shirakawa-mura. Toyama
Prefecture — Mizunashiyama vicinity; Nagato-
okuyama, Fukusawa-mura; Takasugi vicinity,
Ooyama-mura; Onigajo vicinity; Nabemashi vi-
cinity, Shirahagi-mura; Kakumahachi, Matsukura-
mura; Sangayama; Kurobedani; Aimotosanchi vi-
cinity, Aimoto-mura; Ogawadani, Yamazaki-
mura; Taiheisanrin, Sakai-mura. Shizuoka Prefec-
ture— Senzu-goryourin, Kamikawane-mura;
Fujikawayama; Hakkouzan, Shimokawane-mura;
Hakkouzan, Goka-mura; Sekinosawa, Ikawa-
mura; Yakeyama vicinity; Utougi, Ookawauchi-
mura; Yunodake; Uchiyama, Yoshinaga-mura;
Sudoyama vicinity, Sudo-mura; Aitakayama; Kin-
tokiyama; Ita, Heda-mura; Amagisan, Kitou-
mura; Amagisan, Inatori-cho; Amagisan, Kami-
kawatsu-mura; Amagisan, Nishina-mura; Iwashi-
na-mura; Mihama-mura. Nagano Prefecture — Na-
gisoyama, Azuma-mura; Nagisoyama vicinity,
Yomikaki-mura; Tadachi-goryorin; Inagawa-go-
ryorin; Komagatake Mountain Range; Miurayama
vicinity; Mitakeyama-goryorin; Komagane Moun-
tain Range, Hiyoshi-mura; Misogawa-goryorin vi-
cinity, Kiso-mura; Azumi National Forest; Mina-
mikurosawa National Forest, Azusa-mura; Kara-
sugawa National Forest; Karasugawa, Nishihoda-
ka-mura; Kurosawa National Forest, Ogura-mura;
Nakabusa National Forest, Ariake-mura; National
Forest, Matsukawa-mura; National Forest, Toki-
wa-mura; National Forest, Taira-mura; National
Forest, Kamishiro-mura; National Forest, Hokujo-
mura; National Forest, Nakatsuchi-mura; Seto,
Kinasa-mura; Togakushiyama vicinity; Hotokei-
wa, Hirao-mura; Yamada-mura; Shiranesan vicin-
ity; Chichiyama, Toyooka-mura; Nireyama vicin-
ity, Nire-mura; Kakuma National Forest, Osa-
mura; Yatsugatake National Forest, Kitamaki-
mura; National Forest, Minamiaiki-mura;
Yatsugatake National Forest, Minamimaki-mura;
Chichibu Mountain Range; Kamanashiyama; Ni-
shikomagadake, Ina-cho; Fudoutaki; Minamiko-
magadake; Mitsuminegawa, upstream, Inasato-
mura; Akaishisan vicinity, Ooshika-mura; Nishi-
fujiyama, Kami-mura; Kamisawayama, Kisawa-
mura; Nishiyama, Wada-mura; Nishiyama,
Minamiwada-mura; Kumabushiyama; Enasan; Ai-
kawautsubo, Namiai-mura. Niigata Prefecture —
Komagatake vicinity; Koide, Kuramata-mura;
Mukaiyama, Mitsumata-mura; Mantaroyama vi-
cinity, Tsuchidaru-mura; Ushigadake, Ikazawa-
mura; Hakkaisan, Jounai-mura; Hakkaisan, Higa-
shi-mura; Daikokuyama vicinity; Akagawaomote,
Yunotani-mura; Asakusayama vicinity, Irihirose-
mura; Kurohimeyama, Ukawa-mura; Komeyama,
Noda-mura; Kurohimeyama, Takayanagi-mura;
FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES
111
Awagatake; Kawachiyama, Kawachi-mura; Mika-
gura, Nishikawa-mura; Zoukiyama, Higashikawa-
mura; lidesan vicinity, Toyomi-mura; Nagahashiri
vicinity, Hideya-mura; Akataniyama, Akatani-
mura; Ooishifukagawairi vicinity, Sekitani-mura;
Kamikaifu-mura; Komagatake vicinity, Takane-
mura; Wasedasuirin vicinity, Shionomachi-mura;
Fujisawa vicinity, Shimokaifu-mura; Kumadaya-
ma, Nakamata-mura. Yamanashi Prefecture — Ji-
zoutouge vicinity; Tsukiyonodan; Furuyajo, To-
yooka-mura; Takatoriyama; Shichimensanroku,
Motodate-mura; Zarugadake vicinity; Daikoku-
yama; Bettou, Misato-mura; Daitouzan vicinity;
Okusenju, Hirabayashi-mura; Norogawairi, Ashi-
yasu-mura; Kinpusan vicinity, Miyamoto-mura;
Kanegatake vicinity, Kiyokawa-mura; Gozaishi-
yama vicinity, Seitetsu-mura; Gozaishi, Maruno-
mura; Ohsawakanrin, Hourai-mura; Komagatake
Mountain Range; Obi, Masutomi-mura; Namesa-
wayama, Mitomi-mura; Shougenyama; Tokusa.
Hagiwara, Oofuji-mura; Ushiokuyama vicinity,
Okunoda-mura; Ootsuneki, Tabayama-mura; Ko-
ganezawa vicinity, Nanaho-mura; Saihara-mura;
Shimohatsugari, Hatsugari-mura; Okuno, Sasago-
mura; Takara-mura; Mitsutouge, Nishikatsura-
mura; Doushi-mura; Junigatouge. Nagahama-
mura; Oudake; Amegatake; Tojiroyama; Tsuba-
kiyama vicinity, Ookouchi-mura; Sano-goryorin,
Sakae-mura. Tokyo Metropolis — Kanotoyama vi-
cinity; Iriokuyama vicinity; Tarusawayama vicin-
ity; Bonhoriyama. Kanagawa Prefecture — Ashi-
gara Mountain Range; Kintokiyama vicinity;
Bukkazan; Kanazawayama vicinity; Hirugadake.
Chiba Prefecture — Takagoyama; Shimizusawa,
Otaki, Toyooka-mura; National Forest vicinity,
Kameyama-mura; Tsutsumori National Forest, Oi-
kawa-mura. Saitama Prefecture — Ryogamisan;
Ooborayama; Hinoyama; Arimayama; Bukouzan,
Kagemori-mura; Bukouzan, Yokoze-mura. Ihar-
aki Prefecture — Tsukubasan. Tochigi Prefecture —
Konan National Forest vicinity, Ashio-machi; Fu-
kazawa National Forest, Nikko-machi; Fujimiya-
ma vicinity; Tsukiyama vicinity; Ogadake; Taka-
harasan vicinity; Oosabiyama vicinity. Gunma
Prefecture — Hondani National Forest vicinity,
Ueno-mura; Hoshio National Forest vicinity, Oza-
wa-mura; Nishinomaki, Saimoku-mura; Myogisan
National Forest; Nakakimura; Maruyama, Ubu-
chi-mura; Manzayama, Tsumagoi-mura; Yokoka-
be National Forest vicinity, Naganohara-mura; Iri-
yamagawaura National Forest, Kuni-mura; Dai-
gentasan; Bunajurin, Minakami-mura; Hotakaya-
ma vicinity, Kawaba-mura; Hotakayama vicinity,
Katashina-mura; Hirakawa vicinity, Azuma-mura;
Akagiyama vicinity, Akagine-mura; Akagiyama,
Kurohone-mura. Fukushima Prefecture — Futama-
tayama, Yumoto-mura; Kasshizan, Nishigou-
mura; Yaheishiroburaku, Okugawa-mura; Jinku-
rousanrin-oku, Hibara-mura; Azumasan, Azuma-
mura; Shinzan, Mizuho-mura; Moniwa-mura;
Ryozen, Ryozen-mura; Asahidake, Yamakiya-
mura; Hiyama, Asahi-mura. Yamagata Prefec-
ture— Azumayama, Minamihara-mura; Nishine,
Kitaokuni-mura; Fukuei-mura; Rokujuri. Hongou-
mura; Rokujuri, Azuma-mura; Shinzan adjacent
to Mogami-gun, Oosawa-mura; Shinzan adjacent
to Mogami-gun, Nikko-mura; Kuromoriyama vi-
cinity, Nishiokuni-mura; Maemoriyama vicinity,
Higashiokuni-mura; Kamitakarazawa, Higashisa-
wa-mura. Miyagi Prefecture — Ryozen, Hippo-
mura; Oouchi-mura; Oofukasawayama, Shichiga-
shuku-mura; Kuraishidake, Miya-mura; Kawane-
dake National Forest vicinity; Daitoudake vicini-
ty, Akiu-mura; Takakurayama, Ohsawa-mura;
Ooshibayama vicinity, Onikoube-mura; Funako-
shi, Jugohama-mura; Makinosaki vicinity, Ooh-
ara-mura. Akita Prefecture — Ookawamae Nation-
al Forest, Funaoka-mura; Iwamiyama National
Forest vicinity, Iwamisannai-mura; Hatta, Taihei-
mura; Shirayama National Forest, Kamishinjou-
mura; Okuyamasawa National Forest, Kamikoani-
mura; Iwase National Forest, Yamase-mura; Ma-
sezawa, Hachimori-mura; Ozawayama, Sawame-
mura; Kasuge-mura; Fujikotozawa National
Forest vicinity. Fujikoto-mura; Towadako Lake
vicinity. Iwate Prefecture — Goyosan, Hikoroichi-
mura; Tsuchikurayama, Kamiarusu-mura; Utou,
Otsukirai-mura; Shobuzaki, Yoshihama-mura;
Kirigatana, Touni-mura; Komagatake, Kanega-
saki-mura; Samusawayama, Oota-mura; Minami-
toyosawayama vicinity, Yukuchi-mura; Wainai,
Kariya-mura; Fuefuki National Forest, Aozasa-
mura; Kataiwa National Forest, Kamigou-mura;
Matsukurayama, Kasshi-mura; Katahayama, Ku-
rihashi-mura; Tsunagi, Yamagata-mura. Aomori
Prefecture — National Forest, Iwasaki-mura; Na-
tional Forest, Fukaura-mura; National Forest,
Oodose-mura; National Forest, Akaishi-mura; Ka-
waratai vicinity, Nishimeya-mura; Tanosawa vi-
cinity, Uchigata-mura; Yotsugadake vicinity, Aiu-
chi-mura; Nakakodomariyama, Kodomari-mura;
Sanyoushi, Minyama-mura; National Forest, Ima-
betsu-mura; National Forest, Tairadate-mura; Ya-
bitsuyama National Forest, Kanita-mura; Azuma-
dake vicinity, Azumadake-mura; Towada, Houo-
kuzawa-mura; Yaichirousawa vicinity, Oohata-
mura; Hachimoriyama, Kazamaura-mura;
112
FIELDIANA: ZOOLOGY
Futamatayama, Oo-oku-mura; Nuidouishiyama;
Gendoushiro National Forest, Wakinosawa-mura.
KINKAZAN (= KINK AS AN): Miyagi Prefec-
ture— Kinkazan vicinity, Ayukawa-mura.
Appendix 3. Locality Records
(Specimens Examined and Literature
Records) of Macaca fuscata,
Excluding Localities Recorded in
Questionnaire Surveys Conducted by
K. Hasebe in 1923 (Fig. 2A, Appendix
2), K. Kishida in 1953, and H.
Takeshita in 1964 (see above,
Geographic Distribution and Total
Population Estimate); for
Documentation, see Gazetteer,
Appendix 4.
Key to Locality Numbers
Northeastern section (NE)
1. Oma-machi
2. Sai-mura
3. Okoppe River
4. Bushidomari
5. Kusodomari; Shimokita Peninsula; Wakino-
sawa-mura vicinity
6. Kanita-machi
7. Okuradake
8. Ajigasawa-machi
9. Tsugaru-touge
10. Nishimeya-mura
11. Hachimori-machi
12. Yamagata-mura
13. Akita-city
14. Kawabe-machi
15. Nangai-mura
16. Chokai-machi
17. Ogachi-machi
18. Genbikei
19. Sumita-cho
20. Goyosan
21. Ofunato-city
22. Kinkazan, Locs. 1, 2, 6, 10-15, 20, 24, 28,
33, Y6
23. Narusegawa
24. Tagajo-ato
25. Tarogawa
26. Futakuchi
27. Higashine-city
28. Yamadera
29. Kaminoyama-city
30. Kaisho
3 1 . Komogawa; Takane
32.
33.
34.
35.
36.
37.
38.
39.
40.
41.
42.
43.
44.
45.
46.
47.
48.
49.
50.
51.
52.
53.
54.
55.
56.
57.
58.
59.
60.
61.
62.
63.
64.
65.
66.
67.
68.
69.
70.
71.
72.
73.
74.
75.
76.
77.
78.
79.
80.
81.
82.
83.
84.
Asahi-mura
Kurokawa-mura
Takahata-machi
Shichigashuku-machi
Azumayama
Fukushima-city
Soma-city
Kashima-machi
Shiobara-mura
Kawajionsen
Kuriyama-mura
Imaichi-city; Nikko-city
Ashio-machi
Karuizawa-machi
Sanada-machi
Sugadaira
Shiga-kogen (= Shiga Heights)
Jigokudani; Karitayama; Maki; Masugata; Mi-
yamura; Nakayama; Ogikubo; Samusawa;
Shiono; Takai; Tenjinbara; Wakamiya-Hachi-
manjinja; Warabitaira
Wakahowatauchi
Atosugiyama
Asahi-machi
Unazuki-machi
Kurobe
Kamiichi-machi
Oosawano-machi
Hosoiri-mura
Hakusan
Neo-mura
Shichisou-cho
Inuyama-city; Ohirayama
Okinoshima
Oshiro
Shinshiro-city
Hamakita-city
Tatsuyama-mura; Tenryu-city
Toei-cho
Motomura; Suzawa
Kamimachi; Nakadachi
Aoki; Fukagasawa; Irisawai; Kashio; Metaka;
Nakahora; Nishi; Ochiai; Ookawara; Shimizu;
Shimoichiba
Buna; Katsurashima; Yanagisawa
Horane
Kamiseinaiji; Kawasuso; Shimoseinaiji; Taki-
nozawa
Azumabashi
Kakisore
Kamidaira; Kuwasononanbu; Miyamoto; Mi-
yashita; Oojimasan; Tazawa; Tsukiyodaira;
Ushimaki; Yamabuki
Kasugadaira; Kitamura; Shinyashiki
Higashiina
Nishiharuchika
Higashino; Kiso
Oobara; Shouzawa
Hinata
Outaki-mura; Seto, Outaki-mura
Takigoshi
FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES
113
130°
135°
85. Furukaidou; Konishi
86. Fukasawa; Kurouchi; Misokawa
87. Kamikochi
88. Ohamidake; Yarigatake
89. Hida-sanmyaku
90. Hinatayama; Omachi-city; Oosaki; Tokiwa
91. Omachi, hills near
92. Inokamireien; Kawanishi; Miyashiro: Nezumiana
93. Hodaka; Hodaka Golf Course; Tatai
94. Shimashima; Yakeyama
95. Minamimachi: Nakasu
96. Mukawa-mura
97. Kawaguchi-ko
98. Fujiyoshida-city: Nishikatsura-cho
99. Fuji-san
100. Nishiizu-cho
101. Hagachizaki; Ihama
102. Atami-city
114
FIELDIANA: ZOOLOGY
a
Northeastern section (NE)
Specimens examined
Literature records
Introduced populations
Extinct populations
Subfossils
56^55
57
1 92
*393
40°
34 35
43/
84.83828180
^>7
75. 77J
, <• f 71
m
VJ°9o8o107
• ~ 99 1045
103%/
,67
rf^
Q
140°
35°
FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES
115
103.
104.
105.
106.
107.
108.
109.
110.
111.
112.
113.
114.
115.
116.
117.
118.
119.
120.
121.
122.
123.
124.
125.
126.
127.
128.
129.
Hakone-machi; Yugawara-machi
Odawara-city
Ninomiya-machi
Isehara-city
Atsugi-city
Aikawa-machi; Kiyokawa-mura; Tsukui-machi
Tanzawayama
Fujino-machi; Takaosan
Hachioji-city
Hinohara-mura; Itsukaichi
Ome-city
Tabayama-mura
Okutama-machi
Arakawa-mura
Ashigakubo; Yokoze-machi
Houtosan
Kimitsu-city; Otsubo
Aikawa, Futtsu-city; Boso Peninsula (ca.
35°16'N); Futairi; Futtsu-city; Hagiu; Hinaba-
take; Hirooka; Hongou; Kiwadahata; Kyonan-
machi; Nishihikasa; Nutazawa; Okugome;
Okuyama, Kyonan-machi; Ooiwa; Shigumi;
Shimosakuma; Shippara; Tagura; Takagoyama
A; Takagoyama S; Takagoyama Tl; Yamanaka
Yonezawa
Mera
Shirahama-machi
Boso Peninsula, (ca. 35°00'N).
Fukurogura; Kitakomachi; Kiyosumiyama;
Okutani; Sunada; Utsutsumi
Kamou; Kokurano; Kozawamata; Kuradama;
Ootashiro; Tashiro; Tsukagoshi; Tsutsumori;
Yokose
Ishizuka; Koshikiya; Tokuji; Tsukizaki
Kayano-Nanamagari; Sanda
Mobara-city
Southwestern section (SW)
1.
Mikata-cho
2.
Mihama-cho, Fukui
3.
Makino-cho
4.
Nishiazai-cho
5.
Yogo-cho
6.
Kohoku-cho; Takatsuki-cho
7.
Ibuki-cho
8.
Ryozenyama
9.
Taga-cho
10.
Hatasho-cho
11.
Eigenji-cho
12.
Hokusei-cho
13.
Inabe-cho
14.
Yunoyama, Komono-cho
15.
Tsuchiyama-cho [1];
Tsuchiyama-cho [2]
16.
Hino-cho
17.
Koka-cho
18.
Kameyama-city
19.
Oyamada-mura
20. Iga-cho
21. Muroji
22. Oomiya-cho
23. Gokashowan; Nansei-cho
24. Owa.se-city
25. Kumano-city
26. Mihama-cho, Mie
27. Hongu-cho
28. Kozagawa-cho
29. Tsubaki Wild Monkey Park
30. Shirahama-cho
31. Kanaya-cho
32. Tomogashima
33. Minoo-city
34. Arashiyama
35. Otsu-city
36. Hieizan
37. Kamigamo
38. Hiyoshi-cho
39. Mizuho-cho; Sasayama-cho
40. Kasuga-cho
41. Wachi-cho
42. Doumoto; Kidani; Naka; Natasho-mura
43. Adogawa-cho
44. Kaminaka-cho
45. Hotokedani
46. Takahama
47. Otoumi
48. Ine-cho
49. Takeno-cho
50. Wakasa-cho
51. Funakoshiyama
52. Kumayama-cho
53. Kanbanotaki
54. Katsuyama-cho
55. Gagyusan
56. Tenjinkyo
57. Enmeikyo
58. Taishakukyo
59. Akashi; Fukano; Higashihinobori; Houin;
Kisuki-cho; Koujiro; Mitoya-cho; Nishihi-
nobori; Okuyama, Mitoya-cho; Takuwa;
Tsukinoya; Yoshida-mura
60. Ooda-city
61. Mikusu
62. Hasumi-mura; Hinagawa; Hirasa; Imanishi;
Shimogou; Ueda; Ushiroyama
63. Kochi-cho
64. Mihara-city
65. Miyajima-cho
66. Akiyoshi Cave
67. Omishima
68. Choshikei; Rosando; Shodoshima [1]; Sho-
doshima [2]
116
FIELDIANA: ZOOLOGY
69. Awajishima; Kaminada
70. Jinrio (= Jinryo)
71. Anan-city
72. Tachibana
73. Kushigawa
74. Aki-city
75. Chikaraishi
76. Hiromi-cho
77. Nametoko
78. Matsuno-cho
79. Nakamura-city
80. Odo; Odomisaki; Okuuchi-mura
81. Kashima, Ehime
82. Kamae-machi
83. Takasakiyama
84. Kawaradake
85. Gongenyama
86. Hiradoshima
87. Kyogatake; Tara-dake
88. Kugino-mura
89. Higo
90. Itsuki-mura
91. Sagara-mura
92. Kanayama
93. Satsuma-cho
94. Ichiki-cho
95. Kanoya-city
96. Kushima-city
97. Kojima; Torishima
98. Toi-misaki
99. Tanegashima
100. Issou; Kamiyaku-cho [1]; Kusugawa; Mi-
yanoura; Shirakoyama; Yakushima, Loc. P;
Yakushima, Loc. Q
101. Hanyama; Kamiyaku-cho [2]; Kannonzaki;
Kawara; Kojiba; Kuniwaridake, Kamiyaku-
cho; Kuniwaridake, southeastern slopes;
Kuniwaridake, summit area; Nagata; Naga-
ta-todai; Nina-A; Seibu-rindou; Shikamiza-
wa; Yaku vicinity; Yakushima, Loc. A-Loc.
O; Yakushima, southwestern; Yakushima,
western, road vicinity, high zone; Yakushi-
ma, western, road vicinity, low zone; Yaku-
shima, western, road vicinity, middle zone
102. Hinokuchi, Yaku-cho; Koshima; Mugio;
Nakama; Nakase; Nakasegawa; Shichigo-
dake; Yakushima, Loc. R-Loc. U; Yudomari
FOODEN AND AIML SYSTEMATIC REVIEW OF JAPANESE MACAQUES
117
Appendix 4: Gazetteer of Macaca fuscata
Collection and Observation Localities
The sequence of information presented in this
gazetteer is as follows:
1. LocaHty name, in roman letters and
Kanji/Kana characters.
2. Name of prefecture (italics) and island (capi-
tal letters).
3. Coordinates of locality.
4. Date of collection or observation.
5. Name of collector or observer.
6. Bibliographic reference (in parentheses) to
published or unpublished field notes, if any,
7. Abbreviated name of museum (see Introduc-
tion) where specimens are preserved.
8. Number of specimens available (with indi-
cation of part preserved, if skin and skull are
not both present).
9. Locality number (italicized) as indicated in
distribution maps (Appendix 3); this does
not apply to Hasebe's 1923 survey records,
which are mapped in Figure 2A.
Abeyama, Naka-son, Kawakami-gun (JI|_t?P4'
WPrI^Oj); Okayama, HONSHU; 34°48'N,
133°29'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 27: XV-2). See Fig. 2A. ^
Adogawa-cho, Takashima-gun (iliS?P^^JI|
mi); Shiga, HONSHU; ca. 35°19'N, 136°01'E;
mtDNA samples collected before 2003 by Y.
Kawamoto (2002, p. 60). SW43.
Aikawa, Futtsu-city ( S /$ T^ ffi Jll ); Chiba,
HONSHU; 35°11'N, 139°53'E; collected 18
Dec. 1998-30 Mar. 1999 by PRIKU staff;
PRIKU, 5 (skeletons only). NE120.
Aikawa-machi, Aikou-gun ( S ¥ ?P ^ Jl| fflJ );
Kanagawa, HONSHU; ca. 35°29'N, I39°17'E;
mtDNA samples collected before 1998 by Y.
Kawamoto (1997, p. 33). NEWS.
Aikawautsubo, Namiai-mura, Shimoina-gun (T
^Mmi^^¥i^ )\\0'y -^^y, Nagano,
HONSHU; 35°22'N, 137°41'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 47: XXXI-53). See
Fig. 2A.
Aikawayama, Misho-cho, Minamiuwa-gun (1^^
^?P=!®SfflI*IJIiai); Ehime, SHIKOKU;
32°59'N, 132°34'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 18: IX-2). See Fig. 2A.
Aimotosanchi vicinity, Aimoto-mura, Shimonii-
kawa-gun (Tiff JH IPM^^^^ Ol Hfe^);
Toyama, HONSHU; 36°52'N, 137°35'E; re-
ported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 42:
XXIX-10). SeeFig. 2A.
Aitakayama, Takane-mura, Sunto-gun (^^^
fll^^SliUJ); Shizuoka, HONSHU; 35°09'N,
138°49'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p.45:XXX-14). SeeFig.2A.
Ajigasawa-machi, Nishitsugaru-gun (H/^fe?P
0<7-)Rarr); Aomori, HONSHU; ca. 40°47'N,
140°12'E; mtDNA sample collected before
1998 by Y. Kawamoto (1997, p. 33; 2002, p.
60). NE8.
Akagawaomote, Yunotani-mura, Kitauonuma-
gun (db^)S?P^;^^^#JI|^); Niigata,
HONSHU; 37°11'N, 139°03'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 47: XXXII-9). See
Fig. 2A.
Akagiyama, Kurohone-mura, Seta-gun {^^W>
H ft ^ ^ # ^ UJ ); Gunma, HONSHU;
36°3rN, 139°16'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 52: XL- 17). See Fig. 2A.
Akagiyama vicinity, Akagine-mura, Tone-gun
( ^IJ a IP # ^ a ^ '* ^ OJ ffe ); Gunma,
HONSHU; 36°35'N, 139°13'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 52: XL- 16). See Fig.
2A.
Akago, Kitayoshii-mura, Onsen-gun (/m^lPdb
B#W'^^^); Ehime, SHIKOKU; 33°50'N,
132°56'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 21:IX-24). SeeFig. 2A.
Akaioku, Nunuki-mura, Hikami-gun (7KJllP>iS
a:^##^); Hyogo, HONSHU; 35°08'N,
135°00'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 30: XVIII-25). See Fig. 2A.
Akaishisan vicinity, Ooshika-mura, Shi-
moina-gun (T#IP^A^^#^ajfte); Na-
gano, HONSHU; 35°33'N, 138°06'E; reported
FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES
119
in questionnaire suney conducted in 1923 by
K. Hasebe (Iwano. 1974, p. 47: XXXI-46). See
Fig. 2A.
Akaiwa Mountain Range, Maruyae-mura,
Kasa-gun (iPfelPAA>I^#^Ujfl^): Kyoto,
HONSHU; coordinates unknown; reported in
questionnaire sur\ey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 35: XXII-9). Not
mapped.
Akaiwayama, Okadanaka-mura, Kasa-gun (iP-te
iP fS^^ffl 4^ ^ # ^ UJ ); Kyoto, HONSHU;
35°27'N, 135°15'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 35: XXII-8). See Fig. 2 A.
Akakuzureyama. Shimonoho-mura, Mu2i-2un
HONSHU; 35°33'N, 136°59'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 41: XXVIII- 19). See
Fig. 2A.
Akamatsu, Akagaw achi-son, Kaifu-gun (^nP?P
# )! 1*1 ^ # V^ ); Tokushima. ^SHIKOKU;
33°44'N. 134°27'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 23: XI-8). See Fig. 2 A.
Akanishi vicinity, Okutani-mura. Shisou-gun (5^
^lPa^^#ffi^); Hyogo, HONSHU;
35°13'N, 134°3rE; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 30: XVIII-8). See Fig. 2 A.
Akashi, Mitoya-cho, lishi-gun (IS51PH TJMfflJ
a^ S ); Shimane. HONSHU; 35°14'N,
132°50'E; collected 25 Jul 1995 by PRIKU
staff; PRIKU, 1 (skeleton only). SW59.
Akasugiha vicinity, Yamakami-mura, Kan-
zaki-gun (#if AOJ ±^#^!^^itfe); Shiga,
HONSHU; 35°04'N, 136°20'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 38: XXIV-24). See
Fig. 2A.
Akataniyama, Akatani-mura, Kitakanbara-gun
(db«'jiiP#^*\r##aj); Migata, HONSHU;
37°50'N, 139°23'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 47: XXXII-20). See Fig. 2A.
Akataniyama, Ootou-mura, Yoshino-gun ("a©
lPA^^#^aj); Nara. HONSHU; 34°09'N,
135°46'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 35:XXI-2). SeeFig. 2A.
Aki-cit>' (^STtT); KochL SHIKOKU; 33°30'N,
133°54'E; reported present before 1830 by P. F.
von Siebold (MS., 1830-1842, p. I; cf. Tem-
minck. 1842. p. 10; Kuroda, 1938, p. 112;
1940, p. 270). SW74.
Akita-city {^X.^1^): Akita. HONSHU; 39°43'N,
140°07'E; mtDNA extracted from hunting tro-
phy before 2003 by T. Agatsuma and M. Ishi-
gami (2002, p. 79). NEIS^
Akiyoshi Cave. Shuhou-cho, Mine-gun (II^\1P
4li^ ^ ^K^ M ); YamagiichL HONSHU;
34°14'N, 131°18'E; subfossils, possibly older
than Early Jomon (possibly >12 Ka), reported
by M. Kuroda (2002a, p. 115); Kyoto Univer-
sity Anthropology Laboratory. 2 (skulls only,
not seen). SW66.
Akoumayama. Toyonaga-son. Atetsu-gun (P5f|^
IP M ^ ^ # .1 UJ V rt ^ 111 ); Okayama,
HONSHU; 34°58'N. 133°35'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano. 1974. p. 27: XV-12). See Fig.
2A.
Akutayama. Tsugayuki-mura, Oochi-gun (e^
?P tP M ^ ^^^ \h ); Shimane. HONSHU;
35°00'N, 132°38'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano. 1974. p. 28: XVI-24). See Fig. 2A.
Amagisan, Inatori-cho, Kamo-gun (Mj^lPHUi
IHI^i^aj); Shizuoka, HONSHU; 34°46'N,
139°03'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano. 1974,
p. 45: XXX- 19). SeeFig. 2 A.
Amagisan. Kamikavvatsu-mura, Kamo-gun (l£^
iP±)p[^^;f^t^^UJ); Shizuoka, HONSHU;
34°49'N, 138°56'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 45: XXX-20). See Fig. 2 A.
Amagisan. Kitou-mura. Kamo-gun (MS?P^^
:H"^^^aj); Shizuoka. HONSHU; 34°50'N,
139°04'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p.45:XXX-18). SeeFig.2A.
Amagisan, Nishina-mura, Kamo-gun (SM^Pil-
^i^t^^LiJ); Shizuoka, HONSHU; 34°47'N,
138°46'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p.45:XXX-21). SeeFig. 2A.
Amagoitaki waterfall vicinity. Shinritsu-mura,
120
FIELDIANA: ZOOLOGY
Uma-gun (^JSl5|/f II;f^)■M'£)tftfe); Ehime,
SHIKOKU; 33°56'N, 133°40'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 18: IX- 12). See Fig.
2A. _
Ambo-Onoaida( ^M—M'i^f^). See Yakushima,
Census Area 4.
Amegadake, Furuseki-mura, Nishiyatsushiro-gun
(ffi A ft ^ * H ^ M <r S ); Yamanashi,
HONSHU; 35°28'N, 138°32'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 49: XXXIII-34). See
Fig. 2A.
Anan-city (PRjUTtT); Tokiishima, SHIKOKU; ca.
33°55'N, 134°39' E; mtDNA samples collected
before 2003 by Y. Kawamoto (2002, p. 60).
SW71.
Anman-dake. See Kyogatake.
Aoki, Ooshika- mura, Shimoina-gun (~F'^i9P^
±^¥im:^)\ Nagano, HONSHU; 35°32'N,
138°02' E; collected 26 Mar. 1998 by M. Aimi;
PRIKU , 2 (skeletons only). NE70.
Aomishima. See Omishima.
Aonogou-mura, Ooi-gun (AtS^W^^); Fukui,
HONSHU; 35°28'N, 135°30'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 39: XXV-2). See Fig.
2A.
Arakawa-mura, Chichibu-gun (^5<^^^JI|^);
Saitama, HONSHU; ca. 35°57'N, 139°02'E;
large monkey population in vicinity before
1978 reported by T. Kanaizuka (1977, p. 38).
MtDNA samples collected before 1998 by Y.
Kawamoto (1997, p. 33; 2002, p. 60). NE116.
Arashiyama, Kyoto-city (5?.|[Pr|TMlll); Kyoto,
HONSHU; 35°00'N, 135°41' E; study of group
initiated in 1948, provisoning initiated in 1954
(Huffman, 1991a, p. 23). Birth season,
1957-1966, reported by Kawai et al. (1967. p.
38). Blood samples collected before 1992 by
Nozawa et al. (1991, p. 414; 1996, p. 6).
MtDNA samples analyzed ca. 1986-1991 by
Hayasaka et al. (1991, p. 400). Microsatellite
DNA analyzed before 1996 by Do-
mingo-Roura et al. (1997, p. 358). External
measurements taken before 1997 by Hamada
et al. (1996a, pp. 98, 99). MtDNA samples col-
lected before 2003 by Y. Kawamoto (2002, p.
60). SW34.
Arashiyama vicinity, Matsuo-mura, Kadono-gun
( S i? IP a' M*^ it Lljffe); Kyoto, HONSHU;
34°58'N, 135°40'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 36: XXII-26). See Fig. 2A.
Arashiyama West; Texas, U.S.A.; 28°05'N,
99°15'W; captive group, translocated from
Arashiyama in 1972 (Fedigan, 1991a, p. 56;
Fedigan & Griffin, 1996, p. 375). Not mapped.
Aratani vicinity, Shirakawa-mura, Oono-gun (A
i? fP e Jll ^ .^ ^ fte ); Gifu, HONSHU;
36°13'N, 136°52'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 41: XXVIII-40). See Fig. 2A.
Arida-mura/Higashimuro-gun boundary, Nishi-
muro-gun (ffi^S^PWffl^m^^iP^OjM
fte ); Wakayama, HONSHU; 33°30'N,
135°45'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 34: XX-36). See Fig. 2A.
Ariki National Forest vicinity, Takajo-son, Ki-
tamorokata-gun (dbm^c^Pi^^^i^KS^^
^); Miyazaki, KYUSHU; 3r52'N, 131°11'E;
reported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 15: II-7).
See Fig. 2A.
Arimayama, Urayama-mura, Chichibu-gun (^^
?P M UJ ^ W .1 UJ ); Saitama, HONSHU;
35°54'N, 139°05'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 50: XXXVII-6). See Fig. 2A.
Aritani, Hagiwara-mura, Taki-gun (^'^^i^J^
^ T U ^ y, Mie, HONSHU; 34°22'N,
136°20'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 37: XXIII-27). See Fig. 2A.
Asahi-machi, Shimoniikawa-gun (TI/fJH^^ S
err ); Toyama, HONSHU; ca. 36°57'N,
137°35'E; mtDNA samples collected before
2003 by Y. Kawamoto (2002, p. 60). NE52.
Asahi-mura, Iwafune-gun (^^p?P^S^); Nii-
gata, HONSHU; ca. 38°16'N, 139°32'E; re-
ported present before 1978 by Study Group on
the Present Status of Japanese Monkeys
(1977b, p. 23). MtDNA samples collected be-
fore 2003 by Y. Kawamoto (2002, p. 60).
NE32.
Asahidake, Yamakiya-mura, Adachi-gun (^jM
WiU^^M^M^y, Fukushima, HONSHU;
FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES
121
37°36'N, 140°39'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 52: XLI-12). See Fig. 2A.
Asakusayama vicinity, Irihirose-mura, Ki-
tauonuma-gun {itMi.y^W>X!tMW^M-[hi^y,
Niigata, HONSHU; 37°22'N, 139°05'E; re-
ported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 47:
XXXII-10). SeeFig. 2A.
Asan Mountain Range, Tawa-mura, Ookawa-gun
( A Jll IP ^ ^ ^ PrI M LiJ M ); Kagawa,
SHIKOKU; 34°10'N, 134°11'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 24: XII-4). See Fig.
2A.
Asaoki, Mifune-mura, Minamimuro-gun (^#ft
^^^^^'^Wy, Mie, HONSHU; 33°44'N,
I35°58'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 38: XXIII-57). See Fig. 2A.
Ashigakubo, Yokoze-machi, Chichibu-gun (^;^
^^M^P^X^y, Saitama, HONSHU; ca.
35°58'N, 139°08'E; mtDNA samples collected
before 2003 by Y. Kawamoto (2002, p. 60).
NE117.
Ashigara Mountain Range, Miyagino-mura,
Ashigarashimo-gun (£ffi"F?P^^i?^S:M
111 M ); Kanagawa, HONSHU; 35°16'N,
139°05'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 49: XXXV-3). See Fig. 2A.
Ashimi-mura, Oono-gun (A©?P^^^); Fukui,
HONSHU; 36°03'N, 136°25'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 39: XXV- 19). See Fig.
2A.
Ashio-machi, Kamitsuga-gun (Ji^MlPSMffll);
Tochigi, HONSHU; 36°38'N, 139°27'E; col-
lected 20 Aug. 1986 by TPM staff; TPM, 1
(not seen, data from specimen list). NE44.
Ashiu vicinity, Chii-mura, Kitakuwada-gun (db
^ffllPife^^^r^^ffe); Kyoto, HONSHU;
35°18'N, 135°40'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 36: XXII- 17). See Fig. 2A.
Ashiyama vicinity, Ikuma-mura, Nishimuro-gun
(H^«?P^.l^Fajffe); Wakayama,
HONSHU; 33°41'N, 135°27'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 34: XX-24). See Fig.
2A.
Atagoyama vicinity, Sachiyo-mura, Hikami-gun
(7K±?P^tM:WS^Ujfte); Hyogo, HONSHU;
35°irN, 135°03'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 30: XVIII-24). See Fig. 2A.
Atami-city (l^^rfT); Shizuoka, HONSHU; ca.
35°05'N, 139°04'E; mtDNA samples collected
before 1998 by Y. Kawamoto (1997, p. 33;
2002, p. 60). NE102.
Atosugiyama, Otani-mura, Kitaazumi-gun (db$
*iP'h^W^^^Uj); Nagano, HONSHU;
36°52'N, 137°54'E; collected 14 Jan. 1998 by
M. Aimi; PRIKU, 1 (skeleton only). NE51.
Atsugi-city (jpT^ r{]); Kanagawa, HONSHU; ca.
35°27'N, 139°22'E; mtDNA samples collected
before 1998 by Y. Kawamoto (1997, p. 33).
NE107.
Awagatake, Morimachi-mura, Minamikan-
bara-gun {^MW-^W>^^ ¥iMT Uy Niigata,
HONSHU; 37°28'N, 139°09'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 47: XXXIl-14). See
Fig. 2A.
Awajishima C^^^y Hyogo, AWAJISHIMA;
ca. 34° 14' N, 134°52'E; provisioning initiated
in 1967 (Shidei et al., 1981, p. 18). Blood
samples collected before 1992 by Nozawa et al.
(1991, p. 414; 1996, p. 6). Microsatellite DNA
analyzed before 1996 by Domingo-Roura et al.
(1997, p. 358). External measurements taken
before 1997 by Hamada et al. (1996a, pp. 98,
99). MtDNA samples collected before 2003 by
Y. Kawamoto (2002, p. 60). SW69.
Awano, Kawamata-mura, linan-gun (tSl^fPJII
^ ^ ^ S ); Mie, HONSHU; 34°26'N,
136°18'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 37: XXIII-23). See Fig. 2A.
Ayuhanasanrin, Nishida-mura, Mikata-gun (H^
^'mm^TJ.)\-i-\h^y Fukui, HONSHU;
35°32'N, 135°50'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 39: XXV- 12). See Fig. 2A.
Azumabashi, Nagiso-machi, Kiso-gun (^KelP
^TK^fflJl^SM); Nagano, HONSHU;
35°35'N, I37°35'E; collected 28 Mar. 1998 by
M. Aimi; PRIKU, 1 (skeleton only). NE74.
Ill
FIELD! AN A: ZOOLOGY
Azumadake vicinity, Azumadake-mura, Higa-
shitsugaru-gun (K/$^15:^^:H"K^ftfe);
Aomori, HONSHU); 40°50'N, 140°54'E; re-
ported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 57:
XLVI-17). SeeFig. 2A.
Azumasan, Azuma-mura, Yama-gun (?P^1Pr-
SWl^gllj); Fukushima, HONSHU; 37°37'N,
140°irE; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 52: XLI-8). See Fig. 2A.
Azumayama (^SUJ); Yamagata, HONSHU;
37°44'N, 140°12'E; capture of albino monkey
13 Mar. 1938 reported by N. Kuroda (1940, p.
21\).NE26.
Azumayama, Minamihara-mura, Minamioki-
tama-gun (j^Sl^lPl^J^^^SUj); Yama-
gata, HONSHU; 37°48'N, 140°08'E; reported
in questionnaire survey conducted in 1923 by
K. Hasebe (Iwano, 1974, p. 55: XLII-2). See
Fig. 2A.
Azumi National Forest, Azumi-mura, Minamia-
zumi-gun (^^ftl^^ft^^^ftBW^);
Nagano, HONSHU; 36°13'N, 137°42'E; re-
ported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 46:
XXXI-12). SeeFig. 2A.
Beaverton; Oregon, U.S.A.; 45°29'N, 122°48'W;
captive provisioned group translocated from
Mihara ca. 1965 (Van Horn, 1980, p. 198;
Rostal et al., 1986, p. 453). Not mapped.
Benigadake vicinity, Soni-mura, Uda-gun (^PS
W>^M^%LT ^^); Nam, HONSHU;
34°3rN, 136°08'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 35: XXI-9). See Fig. 2 A.
Bettou, Misato-mura, Minamikoma-gun (I^^JS
IP H M ^^t Sy ^ ); Yamanashi, HONSHU;
35°30'N, 138°20'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 48: XXXIII-8). See Fig. 2A.
Bonhoriyama, Togura-mura, Nishitama-gun (S
^SiPP^^^^iSUj); Tokyo Metropolis,
HONSHU; coordinates unknown; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 49: XXXIV-4). Not
mapped.
Bono-misaki, Nishiminamikata-mura, Ka-
wanabe-gun ( Jll ffl^ffi l^7^^±& y ffllllt^);
Kagoshima, KYUSHU; 31°20'N, 130°14'E;
reported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 12: 1-4).
See Fig. 2A.
Boso Peninsula (M-lvt^ft); Chiba, HONSHU;
ca. 35°00'N, 139°55'E; observed Jan.- Dec.
1997 by K. Aizawa and M. Hagiwara (2001, p.
6). NE124.
Boso Peninsula (M,1vg¥ft); Chiha; HONSHU;
ca. 35°16'N, 139°59'E; field studies conducted
in 1972-1973 reported by K. Fukuda (1975, p.
386) and T. Iwano (1977, p. 46; cf. Study
Group on the Present Status of Japanese Mon-
keys, 1977b, p. 26). MtDNA samples collected
before 2003 by Y. Kawamoto (2002, p. 60).
NE120.
Bukkazan, Susugaya-mura, Aikou-gun (^^IP
^ ^ # ^ ^A ^ Oj ); Kanagawa, HONSHU;
35°3rN, 139°15'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 50: XXXV-6). See Fig. 2A.
Bukouzan, Kagemori-mura, Chichibu-gun (^;5<^
?P ^ ^ ^^ E ¥ liJ ); Saitama, HONSHU;
35°58'N, 139°05'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 50: XXXVII-7). See Fig. 2A.
Bukouzan, Yokoze-mura, Chichibu-gun (l>^^?P
#)i^^E¥UJ); Saitama, HONSHU; 35°58'N,
139°08'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 50: XXXVII-8). See Fig. 2A.
Buna, Matsukawa-machi, Shimoina-gun (T'^IP
IP^iJIIWJE^); Nagano, HONSHU; 35°36'N,
137°57'E; collected 23-Jan.-25 Feb. 1998 by M.
Aimi; PRIKU, 5 (3 skeletons only, 2 cranial
fragments and postcranials only). NE71.
Bunajurin, Minakami-mura, Tone-gun (^'J^^
yY.h^\l\^^^^); Gunma, HONSHU;
36°49'N, 139°0rE; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 51: XL- 12). See Fig. 2 A.
Bushidomari, Wakinosawa-mura, Shimokita-gun
iTitmJ&m)R^^±^^y, Aomori, HONSHU;
41°12'N, 140°47'E; collected in 1965-1966 by
S. Azuma; JMC, 1 (skull only, found on sea-
shore). NE4.
Buttsujisan, Takasaka-son, Toyota-gun (MBB^
m^^il^M^ihy, Hiroshima, HONSHU;
34°28'N, 133°02'E; reported in questionnaire
FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES
123
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 26: XIV-25). See Fig. 2 A.
Byobuiwa, Kadohara, Nakahara-mura, Ma-
suda-gun (MBa?P4^i^WP^i^JPm^); Gifu,
HONSHU; 35°44'N, 137°13'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 41: XXVIII-35). See
Fig. 2A.
Cape Toi, Kushima. See Toi-misaki, Kushima.
Chaen, None-mura, Aki-gun ($S?Pi?^^l^
S); Kochi, SHIKOKU; 33°30'N, 134°16'E;
reported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 22: X-42).
See Fig. 2A.
Chichibu Mountain Range, Kawakami-mura,
Minamisaku-gun (^fejXlPJI|±^^5<:UjM);
Nagano, HONSHU; 35°56'N, 138°36'E; re-
ported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 46:
XXXI-39). See Fig. 2A.
Chichiyama, Toyooka-mura, Kamitakai-gun (_h
i5#?PfiEW?Lai); Nagano, HONSHU;
36°36'N, 138°25'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 46: XXXI-30). See Fig. 2A.
Chikaraishi, Higashitsuno-mura, Takaoka-gun
{m^^Mni^ti^); Kochi, SHIKOKU;
33°24'N, 133°03' E; collected 10 Aug. 2002 by
M. Aimi; PRIKU, \.SW75.
Chishorodani vicinity, Hanase-mura, Atago-gun
(SS?P^W^»tti:S&#ffe); Kyoto,
HONSHU; 35°09'N, 135°48'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 36: XXII-22). See Fig.
2A.
Choenzan, Honjou-mura, Yosa-gun (%^lP^j£
^ :U^ OJ ); Kyoto, HONSHU; 35°44'N,
135°15'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 35:XXII-3). SeeFig. 2A.
Chokai-machi, Yuri-gun (S^JIPB^^W); Akita,
HONSHU; 39°11'N, 140°I1'E; mtDNA ex-
tracted from hunting trophy before 2003 by T.
Agatsuma and M. Ishigami (2002, p. 79).
NEI6.
Choshigatani, Makino, Nishinosho-mura, Taka-
shima-gun (S S^PBH+^ife^^^ ^ ^);
Shiga, HONSHU; 35°28'N, 136°03'E; reported
in questionnaire survey conducted in 1923 by
K. Hasebe (Iwano, 1974, p. 38: XXIV- 10). See
Fig. 2A.
Choshikei, Tonosho-cho, Shozu-gun {^\^3.^:t.
Ji fflj ^k ^ M ); Kagawa, SHODOSHIMA;
34°3rN, 134°15'E; provisioned group; birth
season, 1958-1966, reported by Kawai et al.
(1967, pp. 37, 38). 5^65.
Daieiyama, Nakahagi-mura, Nii-gun (^M^P't'
i^^A^Uj); Ehime, SHIKOKU; 33°55'N,
133°2rE; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 21:1X-15). SeeFig. 2A.
Daigentasan, Niiharu-mura, Tone-gun (^J1S?P^
>p^A3f^;*;aJ); Gunma, HONSHU; 36°47'N,
138°50'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 51:XL-I1). SeeFig. 2A.
Daikokuyama, Miyakogawa-mura, Mi-
namikoma-gun (^ ^S^PUP Jll ^ A M UJ );
Yamanashi, HONSHU; 35°28'N, 138°20'E;
reported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 48:
XXXIII-7). See Fig. 2A.
Daikokuyama vicinity, Mikuni-mura, Mi-
namiuonuma-gun (^^>SiPHH:H"AMUjflfe);
Niigata, HONSHU; coordinates unknown; re-
ported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 47:
XXXlI-8). Not mapped.
Daitoudake vicinity, Akiu-mura, Natori-gun (=S
IJ^iPfJ^^WA^^fte); Miyagi, HONSHU;
38°17'N, 140°3rE; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 55: XLIII-6). See Fig. 2A.
Daitouzan vicinity, Nishiyama-mura, Mi-
namikoma-gun (^i^S^PHlllWA^Lil'te);
Yamanashi, HONSHU; 35°33'N, 138°20'E;
reported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 48:
XXXIII-9). See Fig. 2A.
Dakeyama, Kiyodake-son, Kounu-gun (¥^?P
m^^^\h); Hiroshima, HONSHU; 34°44'N,
133°10'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 26: XIV-32). See Fig. 2A.
Damasanchi, Ichinose-mura, Nishimuro-gun (M
#ftfPTfTy^;^^^ajitk); Wakayama,
HONSHU; 33°42'N, 135°28'E; reported in
questionnaire survey conducted in 1923 by K.
124
FIELDIANA: ZOOLOGY
Hasebe (Iwano, 1974, p. 33: XX-23). See Fig.
2A.
Doumoto, Natashou-mura, Onyu-gun (^^IP^
fflJ±4^M;z|s^); FukuU HONSHU; ca. 35°23'N,
135°40'E; collected Aug.-Nov. 1994 by
PRIKU staff; PRIKU, 2 (skeletons only).
SW42.
Doushi-mura, Minamitsuru-gun (i^tPlilPJil^
^ ); Yamanashi, HONSHU; 35°32'N,
139°02'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 49: XXXIIl-31). See Fig. 2A.
Doutokoyama, Suzuhari-son, Asa-gun (^fe^P
ip ?I ^ ^ ^ UJ ); Hiroshima, HONSHU;
34°37'N, 132°34'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 26: XIV- 12). See Fig. 2A.
Douyama, Hata, Tajibe-son, Atetsu-gun (P5[^?P
:l^ >p pP ^ ^ M OJ ); Okayama, HONSHU;
34°59'N, 133°33'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 27: XV-9). See Fig. 2A.
Ebi-cho, Minamishitara-gun (l^lx^lP/S^fflJ);
Aichi, HONSHU; 35°01'N, 137°35'E; reported
in questionnaire survey conducted in 1923 by
K. Hasebe (Iwano, 1974, p. 40: XXVII-3). See
Fig. 2A.
Ebitadani vicinity, Ootsuka-mura, Nishi-
muro-gun (S^«?P AtP)Rl^I ti'^^fe);
Wakayama, HONSHU; 33°34'N, 135°35'E;
reported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 34:
XX-32). See Fig. 2A.
Egasayama, Shimo-mura, Izushi-gun (t±JS?PS
§^^t>lSUJ); Hyogo, HONSHU; 35°30'N,
135°02'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 30: XVIII-22). See Fig. 2A.
Egawa vicinity, Kamiakizuki-mura, Asakura-gun
( ^ 5t ^ ± 1^ >^ ^ >I Jll ffe ); Fukiwka,
KYUSHU; 33°41'N, 130°50'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 17: Vll-3). See Fig.
2A.
Eigenji-cho, Kanzaki-gun ( tt llif IP ^ /Jl ^ BJ );
Shiga, HONSHU; ca. 35°05'N, 136° 18' E;
mtDNA samples collected before 2003 by Y.
Kawamoto (2002, p. 60). SWII.
Enasan, Chisato-mura, Shimoina-gun (~F'^3P^
^S^SSPOJ); Nagano, HONSHU; 35°26'N,
137°42'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 47: XXXI-52). See Fig. 2A.
Endani vicinity, Kawazoe-mura, Nishimuro-gun
(®#S?PJI|^WR^ffe); Wakayama,
HONSHU; 33°38'N, 135°32'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 34: XX-28). See Fig.
2A.
Enmeikyo, Yamano-cho, Fukuyama-city (^ ill
TtT Ol i? ffll M II l^^ ); Hiroshima, HONSHU;
34°42', 133°22'E; mtDNA samples collected
1998-2000 by I. Yoshimi and H. Takaski
(2003, p. 1\).SW57.
Figo. See Higo.
Fudoutaki, Nishikomagadake, Akaho-mura,
Kamiina-gun (±^3P?P#B^M|6l<rS^il)
>S); Nagano, HONSHU; 35°45'N, 137°58'E;
reported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 46:
XXXl-42). See Fig. 2A.
Fuefuki National Forest, Aozasa-mura, Kami-
hei-gun (Jim^mm^^^^m^^y, Iwate,
HONSHU; 39°20'N, 141°40'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Mito, 1989, p. 25). See Fig. 2A.
Fuji-san (Silll) (= Fuji-yama); Kanagawa,
HONSHU; 35°22'N, 138°44'E; collected be-
fore 1888 by M. Seller; MNHN, 3 (including 2
with skulls in skins). NE99.
Fujigaishi vicinity, Kamo-mura, Nii-gun (Iff M^
iP j^ W ^ <r ^5 -te ); Ehime, SHIKOKU;
33°49'N, 133°13'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 21: IX- 17). See Fig. 2 A.
Fujihashi-mura/Higashimuro-gun boundary, Ni-
shimuro-gun (S#g?Ps— M^^K#i?P#
OjMffe); Wakayama, HONSHU; coordinates
unknown; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 34: XX-37). Not mapped.
Fujikawa vicinity, Tsudai-mura, Hata-gun {^^
IP /$ A ^ K Jll ffe ); Kochi, SHIKOKU;
33°08'N, 132°48'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 21: X-12). See Fig. 2A.
Fujikawayama, Nakakawane-mura, Haibara-gun
( ^ J^ iP 4^ Jll ^ ^ S Jll Ol ); Shizuoka,
FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES
125
HONSHU: 35°05'N. 138°05'E: reported in
questionnaire sur\ey conducted in 1923 by K.
Hasebe (Iwano, 1974. p. 45: XXX-4). See Fig.
2A.
Fujikiya forest. Nanaho-mura. Watarai-gun (i^
^?P-fc^^»>1^Maj^): Mie. HONSHU:
34°23'N. 136°29'E: reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano. 1974. p. 37: XXIIl-32). See Fig. 2A.
Fujikotozawa National Forest vicinity. Fuji-
koto-mura. Yamamoto-gun (lll^fP^^^M
#)RSW^fte): .-lA7/t7r HONSHU: 40=20TsI,
140°47'E: reported in questionnaire survey
conducted in 1 923 by K.. Hasebe ( Iwano. 1 974.
p. 56: XLI V- 12). See Fig. 2 A.
Fujimiyama vicinitv. Kuriyama-mura,
Shio'ya-gun (i^^lPliaj^S±Maj^): 7b-
chigi. HONSHU: 36=52'N. 139°32'E: reported
in questionnaire survey conducted in 1923 by
K. Hasebe (Iwano. 1974. p. 51: XXXIX-4).
See Fig. 2A.
Fujino-machi. Tsukui-gun (^^^IPMSW"):
kanagawa. HONSHU": ca. 35°38'N, 139°irE:
mtDNA samples collected before 1998 by Y.
Kawamoto (1997. p. 33). SEllO.
Fujisawa vicinitv. Shimokaifu-mura, Iwa-
fline-gun (^ISiPT^it^S)R^): Migata.
HONSHU: 38°25'N. 139°3rE: reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano. 1974. p. 48: XXXII-25). See
Fig. 2A.
Fujiyoshida-cit>- ( S ± n^ B9 rfT ); Yamanashi.
HONSHU": ca. 35=29'N. 138°48*E: mtDNA
samples collected before 2003 by Y. Kawa-
moto (2002. p. 60). \E98.
Fukabayama. Suigen-mura. Kikuchi-gun (^^
?P7K)g^)^^Uj): Kumamoro. k\Tj"SHU:
33°00'N. 130°54'E: reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano. 1974. p. 17: IV- 1 1). See Fig. 2 A.
Fukagasawa. Ooshika-mura. Shimoina-2un (T
^m^Am^^^'rn): Xagano. HONSHU:
35°30'N. 138°01'E: collected 14 Sep. 1998 by
M. Aimi: PRIKU. 1 (skeleton only). \E70.
Fukano. Yoshida-mura, lishi-gun (t5^1P"affl
^ ^^ S ): Shimane, HONSHU: 35=1 l-N.
132°55'E: collected 13-15 Dec. 1994 by
PRIKU staff: PRIKU. 2 (skeletons only).
SW59.
Fukasawa, Kiso-mura. Kiso-gun (AelP^fi^
}S)R): Sagano. HONSHU: 35°58' N, 137°46'E;
collected 30 Dec. 1997 and 28 Jan. 1998 by M.
.\imi: PRIKU. 3 (skeletons only). NE86.
Fukashimizu vicinity. Kawakami-mura. Taka-
shima-gun (i^SIPJI|±^;^^7Kffe): Shiga,
HONSHU: 35°25'N. 136°01'E: reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano. 1974. p. 38: XXIV-8). See Fig.
2A.
Fukatani vicinity. Yamanokuchi-mura, Oono-gun
(AgiPaj;^P^}^^ffe): Gifu, HONSHU;
35°59'N. 137°10'E: reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano. 1974. p. 41: XXVIII-37). See Fig. 2 A.
Fukazawa National Forest. Nikko-machi,
Kamitsuga-gun (±l?MiPB3feB0"}^>RSW
^): Tochigi. HONSHU: 36°44'N. 139°35'E;
reported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 51:
XXXIX-3). SeeFig. 2A.
Fukiyama National Forest. Minoo-son,
Kov-u-gun (JPu^^iPZlrt^q^LijSW^); Mi-
xazakL KYUSHU: 32°07'N. 131°19'E: re-
ported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano. 1974. p. 15: 11-17).
See Fig. 2A.
Fukubegadake. Takeda-mura, Gujo-gun (^JilP
* ^^ MS): G/f//, HONSHU; 35°39'N,
136°56'E: reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano. 1974,
p. 41: XXVIII-22). See Fig. 2A.
Fukuei-mura. Nishitagavva-gun (IIBQ JII?P^3^
^): Yamagata. HONSHU: 38°34'N. 139°44'E;
reported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano. 1974, p. 55:
XLII-5). See Fig. 2A.
Fukiihara-son. Katsuura-gun (0/i^?PISi^^);
Tokushima. SHIKOKU: 33°53'N. 134°26'E;
reported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano. 1974. p. 23:
XI-15). SeeFig. 2A.
Fukurogura, Kamogawa-citv- ( fli jll rfT ^ j& );
Chiba, HONSHU: ca. 35°08'N. 140°07'E; col-
lected Sep. 1997 by PRIKU staff: PRIKU, I
(skeleton only). \EI25.
Fukushima-cit> (IS firfJ): Fukushima. HONSHU;
ca. 37°45'N. 140°28'E: blood samples col-
lected before 1992 by Nozawa et al. (1991, p.
126
HELD! AN A: ZOOLOGY
414; 1996, p. 6). MtDNA samples collected
before 2003 by Y. Kawamoto (2002, p. 60).
NE37.
Funakoshi, Jugohama-mura, Monou-gun (^fe^
?P + £ ^ ^^ fa M ); MiyagU HONSHU;
38°30'N, 141°27'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 55: XLIII-10). See Fig. 2A.
Funakoshiyama, Mikawa-mura, Shisou-gun (7^
H ?P H M ;f^t fq^ M liJ ); Hyogo, HONSHU;
35°07'N, 134°25'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 30: XVIII-5). See Fig. 2A.
Funakoshiyama, Nankou-cho, Sayo-gun ("feffl^
]^)tefflTfp'®aj); Hyogo. HONSHU; 35°07'N,
134°25'E; provisioned group; birth season,
1965-1966, reported by Kawai et al. (1967, pp.
37, 38). SW51.
Furukaidou, Kaida-mura, Kiso-gun (T^e^^ffl
^*^^jI); Nagano, HONSHU; 35°58'N,
137°35'E; collected 4 Sep. 1998 by M. Aimi;
PRIKU, 2 (skeletons only). NE85.
Furutani National Forest vicinity, Warabihara,
Kamiosakabe-son, Atetsu-gun (PR[@^±ffJoP
^MJ^*#lW^ffe); Okayama, HONSHU;
35°08'N, 133°32'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 27: XV-8). See Fig. 2A.
Furutayama vicinity, Kitatane-mura,
Kumage-gun (fl^?Pdb«^^* BQ OJ^);
Kagoshima, TANEGASHIMA; 30°44'N,
131°02'E; reported as extinct in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 12: 1-2). See Fig. 2A.
Furuyajo, Toyooka-mura, Minamikoma-gun (1^
^SiPMI^W^^^); Yamanashi,
HONSHU; 35°20'N, 138°24'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 48: XXXIIl-3). See
Fig. 2A.
Futagoyama, Nishimusashi-mura, Higashikuni-
saki-gun (:^S^?PffiEMWi35^Li|); Oita,
KYUSHU; 33°34'N, 13r36'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 16: 111-20). See Fig.
2A.
Futagoyama vicinity, Susami-mura, Nishi-
muro-gun (lS#«lPMl#m^)5(^ UJ ffe);
Wakayama, HONSHU; 33°33'N, 135°32'E;
reported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 34:
XX-30). See Fig. 2A.
Futairi, Kimitsu-city ( ^ /$ rtT ~ A ); Chiba,
HONSHU; 35°13'N, 140°00'E; collected 23
Aug. 1998 by PRIKU staff; PRIKU, 1 (cranial
fragments only). NE120.
Futakuchi, Sendai-city ( fllj n T^ — P ), Miyagi,
HONSHU; 38°16'N, 140°33'E; observed
Jan.-Mar. 2003 by K. Izawa, T. Uno, and H.
Fujita (2003, p. \1).NE26.
Futamatayama, Oo-oku-mura, Shimokita-gun (T
dblPAH^-XUj); Aomori, HONSHU;
41°26'N, 140°56'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 57: XLVI-21). See Fig. 2A.
Futamatayama, Yumoto-mura, Iwase-gun {^Wi.
iPM^^-MliJ); Fukmhima, HONSHU;
37°16'N, 140°00'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 52: XLI-4). See Fig. 2A.
Futtsu-city (s)iTU); Chiba, HONSHU; ca.
35=127^^, 139°53'E; mtDNA samples collected
before 1999 by Y. Kawamoto (1998, p. 54).
NE120.
Gagyusan ( El 4 OJ ); Okayama, HONSHU;
34°49'N, 133°37'E; provisioning initiated in
1955 (Shidei et al., 1981, p. 18). Birth season,
1956-1966, reported by Kawai et al. (1967, p.
39). Liver samples collected from translocated
captives before 1987 by Hayasaka et al. (1986,
p. 346). Blood samples collected before 1992
by Nozawa et al. (1991, p. 414; 1996, p. 6).
MtDNA samples collected before 2003 by Y.
Kawamoto (2002, p. 60). SW55.
Gagyusan, Matsuyama-son, Joubou-gun (_hj^?P
^i^ \h ^ ^ ^ \h ); Okayama, HONSHU;
34°49'N, 133°37'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 27: XV- 17). See Fig. 2A.
Gagyusan, Takahashi-cho, Joubou-gun (JiMlP
i^ ^ fflj ei 4^ Uj ); Okayama, HONSHU;
34°49'N, 133°37'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 27: XV- 16). See Fig. 2A.
Gagyusan, Tsugawa-son, Joubou-gun (_hj^?Py$
JHWEil^Uj); Okayama, HONSHU; 34°49'N,
133°37'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES
127
p. 27: XV- 15). See Fig. 2A.
Gayatani vicinity, Nakatsu-mura, Kamiukena-
gun (±>?7v:^4'^^:^'-\'#^); Ehime.
SHIKOKU; 33°37'N, 133°04'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 21: IX-20). See Fig.
2A.
Genbikei, Genbi-cho, Ichinoseki-city ( — ^TtJJ^
II WJ^ 11)1); hxate. HONSHU; 38°58'N,
141°02'E; mtDNA samples collected before
1998 by Y. Kawamoto (1997, p. 33; 2002, p.
60). NE18.
Gendoushiro National Forest, Wakinosawa-mura.
Shimokita-gun (Tdb^^if)R;^)lK^lW
^); Aomoh, HONSHU; 41°25'N, 140°54'E;
reported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 57:
XLVI-23). See Fig. 2A.
Gezan, Toyotashimo-son, Toyoura-gun (sM?P
S ra T W ^ OJ ); Yamagiichi, HONSHU;
34°10'N, 131°0rE; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 25: XIII-2). See Fig. 2 A.
Ginzan vicinity, Oomori-cho, Nima-gun (SRIP
A^BTr^Ujftfe); Shimane, HONSHU; 35°08'N,
132°28'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 28: XVI-30). See Fig. 2A.
Gokairiaiyama, Kamishou-mura, Oono-gun (A
i^lP±Ji^E'r A^ULl); Fukui. HONSHU;
35°57'N, 136°3rE; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 39: XXV-21). See Fig. 2A.
Gokashowan {^TpfiMy, Mie. HONSHU; ca.
34°20'N, 136°41'E; observed Aug. 1979 by K.
Masui (Takasaki & Masui, 1984, p. 310).
SW23.
Gokurakujisanrin, Hara-son, Saeki-gun (fe'ffilP
ii^S^#lll^); Hiroshima', HONSHU;
34°22'N, 132°20'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 26: XIV- 1). See Fig. 2A.
Gongendaki vicinity. Sakashukitou-son,
Naka-gun (i9m?PJS'>H^IIW=I>y >^ +
^ ); Tokushima, SHIKOKU; 33°50'N,
134°20'E; reported in questionnaire survey
conducted in 1 923 by K. Hasebe (Iwano, 1 974,
p. 23:XI-13). SeeFig. 2A.
Gongenyama, Higashisefuri-son, Kanzaki-gun
(ftif^SW^^^mUj); Saga, KYUSHU;
33°22'N, 130°23'E; reported in 1973 by H.
Ikeda and K. Eguchi (1978, p. 59). SW85.
Gongenyama \icinity, Konose-mura, Kuma-gun
(SSfPtt^^ft^iajfte); Kumamoto,
KYUSHU; 32°18'N, 130°38'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 16: IV-3). See Fig.
2A.
Goriyama. Kawaguchi, Ogawagyou-mura, Wa-
tarai-gun (Jg^lP^hJHiP^JH PSMOj); Mie,
HONSHU; 34°24'N, 136°37'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 37: XXIII-34). See
Fig. 2A.
Gosho-son, Itano-gun (^i?fP1Slp|f^); Toku-
shima, SHIKOKU; 34°09'N, 134°23'E; re-
ported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 24:
XI-35). See Fig. 2A.
Goyosan ( £ ^ Oj ); hvate, HONSHU; ca.
39°12'N, 14r44'E; mtDNA samples collected
before 2003 by Y. Kawamoto (2002, p. 60; cf.
Study Group on the Present Status of Japanese
Monkeys, 1977a, p. \{)).NE20.
Goyosan, Hikoroichi-mura, Kesen-gun (^^fllj^P
B tiltT;f>tE^Uj); hvate, HONSHU; 39°09'N,
141°4rE; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 56:XLV-1). SeeFig. 2A.
Gozaishi, Maruno-mura, Kitakoma-gun (db^0
fPlil©^^^^); Yamanashi, HONSHU;
35°44'N, 138°25'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 48: XXXIII- 15). See Fig. 2A.
Gozaishiyama vicinity. Seitetsu-mura, Kita-
koma-gun (db&0?P^^^^^^Uj^);
Yamanashi, HONSHU; 35°43'N, 138°21'E;
reported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 48:
XXXIII-14). SeeFig. 2A.
Gozenyama, Hagiwara-cho, Masuda-gun (MB9
iPi^J^fflJ^lfUj); Gifu, HONSHU; 35°52'N,
137°16'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p.41:XXVIII-31). SeeFig. 2A.
Hacchoyama, Yuge-mura, Kitakuwada-gun (db
HBaiP^ijtfATOj); Kyoto, HONSHU;
35°12'N, 135°39'E; reported in questionnaire
128
FIELDIANA: ZOOLOGY
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 36: XXII- 18). See Fig. 2 A.
Hachimori-machi, Yamamoto-gun (Lil^t^^PAl^
fflj); Akita, HONSHU; ca. 40°19'N, 140°02'E;
mtDNA samples collected before 1998 by Y.
Kawamoto (1997, p. 33; 2002, p. 60; cf. Study
Group on the Present Status of Japanese Mon-
keys, 1977a, p. 10). Collected before 2002 by
PRIKU staff; PRIKU, 1 (mandible and post-
cranials only). NEl 1.
Hachimoriyama, Kazamaura-mura, Shi-
mokita-gun (TdbiPEPB^/i^A^lij); Ao-
mori, HONSHU; 41°26'N, 141°00'E; reported
in questionnaire survey conducted in 1923 by
K. Hasebe (Iwano, 1974, p. 57: XLVI-20). See
Fig. 2A.
Hachioji-city (Ai-frfT); Tokyo Metropolis,
HONSHU; ca. 35°39'N, 139°20'E; mtDNA
samples collected before 1999 by Y. Kawa-
moto (1998, p. 54). ^£777.
Hachirouyama, Tahara-mura, Higashimuro-gun
( m # « ?P ffl J^ W A HP OJ ); Wakayama,
HONSHU; 33°32'N, 135°54'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 34: XX-43). See Fig.
2A.
Hachisusanrin vicinity, Mori-mura, linan-gun
( IS it IP 1^ ^ a OJ # ); Mie, HONSHU;
34°2rN, 136°08'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 37: XXIII-24). See Fig. 2A.
Hagachizaki, Minamiizu-cho, Kamo-gun (M^
^M^s:fflI):^Blli^) (= Hashozaki); Shizuoka,
HONSHU; 34°41'N, 138°45'E; reported pre-
sent in 1934 by N. Kuroda (1938, p. 1 12; 1940,
p. 270). Provisioning initiated 1953-1955
(Akabori & Suzuki, 1977, p. 73; Study Group
on the Present Status of Japanese Monkeys,
1977b, p. 23; Shidei et al., 1981, p. 17). Birth
season, 1966, reported by Kawai et al. (1967, p.
39). Collected 19 Nov. 1978 by PRIKU staff;
PRIKU, 1 (skeleton only). MtDNA samples
collected ca. 1986-1991 by Hayasaka et al.
(1991, p. 400). Blood samples collected before
1992 by Nozawa et al. (1991, p. 414; 1996, p.
6). Microsatellite DNA analyzed before 1996
by Domingo-Roura et al. (1997, p. 358). Ex-
ternal measurements taken before 1997 by
Hamada et al. (1996a, pp. 98, 99). MtDNA
samples collected before 1998 by Y. Kawa-
moto (1997, p. 33). A^£707.
Hagi-machi, Abu-gun (PRf^lPMBJ); Yamaguchi,
HONSHU; coordinates unknown; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 25: XIII- 10). Not
mapped.
Hagiu, Futtsu-city ( ^ /$ TfT f^ ^ ); Chiba,
HONSHU; 35°10'N, 139°50'E; collected 30
Mar. 1999 by PRIKU staff; PRIKU, 1 (skele-
ton only). NE120.
Hakkaisan, Higashi-mura, Minamiuonuma-gun
(itl^)S^^WA^aj); Niigata, HONSHU;
37°10'N, 138°59'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 47: XXXII-7). See Fig. 2A.
Hakkaisan. Jounai-mura, Minamiuonuma-gun
(^^)SlP^rt^A^^aj); Niigata, HONSHU;
37°08'N, 138°59'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 47: XXXII-6). See Fig. 2 A.
Hakkouzan, Goka-mura, Haibara-gun (^i^?P£
W^e3feUj); Shizuoka, HONSHU; 34°52'N,
138°06'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p.45:XXX-6). SeeFig. 2A.
Hakkouzan, Shimokawane-mura, Haibara-gun
( ^ J^ ?P T Jll ^ ^ e Tfe UJ ); Shizuoka,
HONSHU; 34°58'N, 138°04'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 45: XXX-5). See Fig.
2A.
Hakone-machi, Ashigarashimo-gun (S.ffiT?P^
^ BO" ); Kanagawa, HONSHU; 35°11'N,
139°02'E; provisioning initiated in 1956 (Ta-
naka & Masui, 1977, p. 62). Variable provi-
sioning reported 1971-1977 by F. Fukuda
(1988, p. 480). Intergroup dispersal reported
1966-1980 by F. Fukuda (2004, p. 54).
NE103.
Hakusan (fiOj); Ishikawa, HONSHU; 36°16'N,
136°46'E; field study begun in 1962, provi-
sioning attempted in 1964 (Masui, 1977, p. 81).
Blood samples collected before 1992 by No-
zawa et al. (1991, p. 414; 1996, p. 6). Mi-
crosatellite DNA analyzed before 1996 by
Domingo-Roura et al. (1997, p. 358). External
measurements taken before 1997 by Hamada
et al. (1996a, pp. 98, 99). NE58.
FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES
129
Hamahara-mura, Oochi-gun (e^^P^J^^);
Shimane, HONSHU; 35°03'N, 132°36'E; re-
ported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 28:
XVI-23). See Fig. 2A.
Hamakita-city (^JbTti ); Shizuoka, HONSHU; ca.
34°48'N, 137°48'E; collected before 1991 by
PRIKU staff; PRIKU, 3 (skeletons only).
NE65.
Handairayama-son. See Nagafuka vicinity.
Hanto-cho, Itano-gun (^iJ^P^^fflJ); Tokii-
shima, SHIKOKU; 34°09'N, 134°31'E; re-
ported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 23:
XI-33). See Fig. 2A.
Hanyama, Kamiyaku-cho, Kumage-gun (^^IP
±M:^fflI^Uj); Kagoshima, YAKUSHIMA ;
30°23'N, 130°23'E; collected Mar. 1983 by T.
Maruhashi; PRIKU, 1 (skeleton only). Col-
lected ca. 1983-1999 by PRIKU staff; PRIKU,
30 (skeletons only, including 7 incomplete).
Collected 10 May 1984 by f. Oi; PRIKU, 1
(skull only). Collected Jun.-Sep. 1991 by N.
Agetsuma; PRIKU, 3 (skeletons only). Mass
mortality reported in 1998-1999 by G. Hanya
et al. (2004, p. \^2)SW101.
Hanyama-Segire, Kamiyaku-cho, Kumage-gun
(^^^±M^fflI¥ajM^). See Yaku'^hima,
Census Area 7.
Harimochi, Nishitara-mura, Isa-gun ("^felPM
;*;^^t1-J^); Kagoshima, KYUSHU; 32°02'N,
130°38'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 12:1-9). See Fig. 2A.
Hashozaki. See Hagachizaki.
Hasumi, Hasumi-mura, Oochi-gun (e^?P^^
M^^^H); Shimane, HONSHU; 34°51'N.
132°40'E; collected 6 Dec. 2000 by M. Aimi;
PRIKU, 1 (skeleton only). SW62.
Hata-gun. See Okuuchi-mura.
Hata-mura, lishi-gun (ISS?P/^^W); Shimane,
HONSHU; 35°08'N, 132°39'E; reported (as
possibly temporary habitat) in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 28: XVI-32). See Fig. 2 A.
Hatasho-cho, Echi-gun (^^IP^Sfflf); Shiga,
HONSHU; ca. 35° 10' N, 136°16'E; mtDNA
samples collected before 2003 by Y. Kawa-
moto (2002, p. 60). SWIO.
Hatofukiyama, Dota-mura, Kani-gun (RTJ/^lPih
ffl ^ ii R^ UJ ); Gifu, HONSHU; 35=247^,
137°02'E; reported in questionnaire survey
conducted in 1 923 by K. Hasebe (Iwano, 1 974,
p. 41: XXVIII-21). See Fig. 2A.
Hatta, Taihei-mura, Minamiakita-gun (^^i^ffllP
;l!^¥:^Affl); Akita, HONSHU; 39°45'N,
140°irE; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 56: XLIV-4). See Fig. 2A.
Hattasan vicinity, Todoromi-mura, Toyono-gun
{^m^±^ SM^ABBOJte); Osaka,
HONSHU; 34°53'N, 135°30'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 33: XIX-2). See Fig.
2A.
Hida-sanmyaku (mountain range) (M^lllM);
Nagano, HONSHU; ca. 36°25'N, 137°40'E;
observed before 2004 by J. Ackerman (2003, p.
97). NE89.
Hidaka Mountain Range, Kawakami-mura, Hi-
daka-gun ( S SlPJlfi^ S S UjM); Waka-
yama, HONSHU; 33°58'N, 135°25'E; reported
in questionnaire survey conducted in 1923 by
K. Hasebe (Iwano, 1974, p. 33: XX-9). See Fig.
2A.
Hidarimaeyama vicinity, Tsuji, Momose-mura,
Takashima-gun (l§E^W^Wit:£Bif Ujffe);
Shiga, HONSHU; 35°26'N, 136°03'E; reported
in questionnaire survey conducted in 1923 by
K. Hasebe (Iwano, 1974, p. 38: XXI V-9). See
Fig. 2A.
Hidaritsubute \'icinity. Torigoe-mura, Nomi-gun
{^^^MU^&Mi^y, Ishikawa, HONSHU;
36°18'N, 136°37'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 40: XXVI-4). See Fig. 2A.
Hieizan (J±0Uj); Shiga, HONSHU; ca. 35°05'N,
135°50'E; >12 km troop movement, 26
Oct.-21 Nov. 1996, reported by G. Hanya et al.
(2002, p. 417). MtDNA samples collected be-
fore 2003 by Y. Kawamoto (2002, p. 60).
SW56.
Hieizan. Sakamoto-mura, Shiga-gun (^MIP^
^WJtMOj); Shiga, HONSHU; 35°05'N,
135°5rE; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 38: XXIV-2). See Fig. 2A.
Higakureyama vicinity, Iwato-son, Ni-
130
nELDIANA: ZOOLOGY
shiusuki-gun (ffi Q tt?P^P ^ S H UJ ffe);
Miyazaki, KYUSHU; 32°45'N, 131°25'E; re-
ported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 15: 11-30).
See Fig. 2A.
Higashi- and Nishi-Kurokiyama, Kamiimi-mura,
Higashikunisaki-gun (!^P^?P±^II^^-
S H ?K 111 ); Oita, KYUSHU; 33°37'N,
131°36'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 16:111-21). See Fig. 2A.
Higashihinobori, Kisuki-cho, Oohara-gun (AJ^
^ ^ ;^ fflj K B ^ ); Shimane, HONSHU;
35°16'N, 132°55'E; collected 16 Sep.-22 Dec.
1994 by PRIKU staff; PRIKU, 3 (skeletons
only, 1 without skull). SW59.
Higashiina, Komagane-city {^^^l^Mi^M);
Nagano, HONSHU; 35°45'N, 137°58'E; col-
lected 23 Jul. 1998 by M. Aimi; PRIKU, 1
(skeleton only). NE78.
Higashimikatagamori, Tokuda-mura/Nakagawa-
mura/Sekiya-mura border area, Shusou-
gun(j^^^t§Ba^^Ba)t. ^mmm^tm%
E.y5 r By, Ehime, SHIKOKU; 33°54'N,
132°57'E; reported in questionnaire survey
conducted in 1 923 by K. Hasebe (Iwano, 1 974,
p. 18:IX-9). SeeFig. 2A.
Higashimisaki vicinity, Kuki-mura, Kita-
muro-gun {it^U^%%.^'^%^^^)\ Mie,
HONSHU; 34°01'N, 136°15'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 37: XXIII-47). See
Fig. 2A.
Higashimiyamata vicinity, Kawakami, Ichi-
nose-mura, Watarai-gun (J^^IP — ^i^^^Jil
JiM'BU^); Mie, HONSHU; 34°20'N,
I36°34'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 37: XXIII-35). See Fig. 2A.
Higashine-city (M^'J^); Yamagata, HONSHU;
ca. 38°24'N, 140°25'E; mtDNA samples col-
lected before 2003 by Y. Kawamoto (2002, p.
60). NE27.
Higashino, Agematsu-machi, Kiso-gun {^mW>
±4i^ffll^i?); Nagano, HONSHU; 35°45'N,
137°43'E; collected 25 Mar. and 22 Jun. 1998
by M. Aimi; PRIKU, 5 (skeletons only). NE80.
Higashinokawa vicinity, Oohuki-mura, Nii-gun
(ff^^PA^^^myJIIftfe); Ehime,
SHIKOKU; 33°51'N, 133°11'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 21: IX- 18). See Fig.
2A.
Higashiogura-mura, Echi-gun (M^^^^^J^^^^);
Shiga, HONSHU; 35°06'N, 136°24'E; reported
in questionnaire survey conducted in 1923 by
K. Hasebe (Iwano, 1974, p. 38: XXIV-23). See
Fig. 2A.
Higashiyama National Forest, Yasuharakamin-
ishi-mura, Kagawa-gun (#JI|?P$J^-hM^
^lUm^^y, Kagawa, SHIKOKU; 34°08'N,
134°05'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 24: Xll-3). See Fig. 2A.
Higashiyama vicinity, Kamihaya-mura, Nishi-
muro-gun (H#ft?P±^*^^tKajfte); Wa-
kayama, HONSHU; 33°49'N, 135°25'E; re-
ported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 33:
XX-13). SeeFig. 2A.
Higetayama vicinity, Takane-mura, Masuda-gun
{^m^mm^^f^\hWy, a/u, honshu;
36°06'N, 137°29'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 41: XXVIII-34). See Fig. 2A.
Higo (flE^) (= Figo); Kumamoto, KYUSHU; ca.
32°27'N, 130°46'E; reported present before
1830 by P. F. von Siebold (MS., 1830-1842, p.
1; cf Temminck, 1842, p. 10; Kuroda, 1938, p.
112; 1940,p. 271).5^<^9.
Hikosanroku, Tsukinoki-mura, Shimoge-gun (T
^^P^tK^^^UjM); Oita, KYUSHU;
33°28'N, 130°59'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 16: 111-17). See Fig. 2A.
Hinabatake, Kyonan-machi, Awa-gun (^^?PIS
l^ffllBfSjjffl); Chiba, HONSHU; 35°08'N,
139°52'E; collected 17 Dec. 1997-2 Dec. 1998
by PRIKU staff; PRIKU, 6 (skeletons only).
NE120.
Hinagawa, Hasumi-mura, Oochi-gun (e^^^
MMWS^JII); Shimane, HONSHU; 34°51'N,
132°40'E; collected 18 Jun. 1998-15 Jan. 1999
by PRIKU staff; PRIKU, 3 (skeletons only).
Collected 8 Dec. 2000 by M. Aimi; PRIKU, 1
(skeleton only). SW62.
Hinata, Mitake-mura, Kiso-gun (TKe^PH^^t^
B fS]); Nagano, HONSHU; 35°50'N, 137°39'E;
FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES
131
collected 30 Jul. 1998 by M. Aimi; PRIKU, 1
(skeleton only). NE82.
Hinatayama, Oomachi-city (AffljTfJ B [d]i1|);
Nagano, HONSHU; 36°32'N, 137°47'E; col-
lected 19 Sep. and 19 Dec. 1997 by M. Aimi;
PRIKU, 2 (skeletons only). NE9Q.
Hino-cho, Gamou-gun (S^?PS©fflI); Shiga,
HONSHU; ca. 35°01'N, 136°15'E; mtDNA
samples collected before 2003 by Y. Kawa-
moto (2002, p. 60). SW16.
Hinohara-mura, Nishitama-gun (B^BIP^J^
^); Tokyo Metropolis, HONSHU; ca. 35°43'N,
139°10'E; mtDNA samples collected before
1999 by Y. Kawamoto (1998, p. 54; 2002, p.
6Q).NE112.
Hinokiyama, Toumi-son, Higashiusuki-gun (^
Q^IPK^W^UJ); Miyazaki, KYUSHU;
32°38'N, 131°38'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 15: 11-26). See Fig. 2 A.
Hinokuchi, Yaku-cho, Kumage-gun (^^?PM
:^ HI ffi y P ); Kagoshima, YAKUSHIMA;
30°17'N, 130°38'E; collected 25 Feb. 1989 by
PRIKU staff; PRIKU, 5 (skeletons only, in-
cluding 1 with mismatched mandible). SIV102.
Hinoyama, Shirakawa-mura, Chichibu-gun {^1>t
?P S Jll ^ S © UJ ); Saitaina, HONSHU;
35°56'N, 138°59'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 50: XXXVII-5). See Fig. 2A.
Hippo-mura, Igu-gun (#M^^^^ : Hfil^M
Uj <i; U < ^ ); Miyagi, HONSHU; 37°50'N,
140°44'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 55:XLIII-1). SeeFig. 2A.
Hiradoshima, Kitamatu'ura-gun (db^MlP^P
S); Nagasaki, HIRADOSHIMA; 33°20'N,
129°28'E; observed until ca. 1925 by local
residents; subsequently extinct on this island
(Sakura, 1976, p. \5\). SW86.
Hirai vicinity, Kawakami-son, Kaifu-gun {M^
iPJI|±^¥#^); Tokushima, SHIKOKU;
33°39'N, 134°17'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 23: XI-2). See Fig. 2A.
Hirakawa vicinity, Azuma-mura, Tone-gun (^0
^ IP ^ ^ ¥ Jll ffe ); Gunma, HONSHU;
36°42'N, 139°19'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 51: XL- 15). See Fig. 2A.
Hiraki vicinity, Nagano-mura, Ano-gun ($/SlP
:^i?^¥-^'te); Mie, HONSHU; 34°45'N,
136°2rE; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 36:XXIII-13). SeeFig. 2A.
Hirakura, Yahata-mura, Ichishi-gun ( — ^vlPA
<§ W ¥ )t ); Mie, HONSHU; 34°29'N,
136°15'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 37:XXIII-21). SeeFig. 2A.
Hirasa, Hasumi-mura, Oochi-gun (e^?P^^
ll^¥ft); Shimane, HONSHU; 34°51'N,
132°41'E; collected 23 Jul.-2 Dec. 1995 by
PRIKU staff; PRIKU, 5 (skeletons only, in-
cluding 1 without skull). SW62.
Hirasan vicinity, Katsuragawa-mura, Shiga-gun
(^^M?P«Jll^l±maj^); Shiga, HONSHU;
35°14'N, 135°52'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 38: XXIV-4). See Fig. 2A.
Hirasan vicinity, Kido-mura, Shiga-gun (^MIP
TKP^hbMUjfe); Shiga, HONSHU; 35°12'N,
135°55'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 38: XXIV-3). See Fig. 2A.
Hiromi-cho, Kitauwa-gun (db^^lPJABBTT);
Ehime, SHIKOKU; ca. 33°15'N, 132°42'E;
mtDNA samples collected before 2003 by Y.
Kawamoto (2002, p. 60). SW76.
Hirooka, Kimitsu-city ( ^ /^ Tfi" JA [55 ); Chiba,
HONSHU; 35°15'N, 140°03'E; collected 14
Nov. 1997 by PRIKU staff; PRIKU, 1 (skele-
ton only). NE120.
Hirosesanrin, Tenkawa-mura, Yoshino-gun (^
S^P^JIIWJA^OJ^); Nara, HONSHU;
34°14'N, 135°52'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 35: XXI-3). See Fig. 2 A.
Hirugadake, Toya-mura, Tsukui-gun (^A.^^
1 Jl ^ 4^ if' U ); Kanagawa, HONSHU;
35°3rN, 139°12'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 50: XXXV-8). See Fig. 2A.
Hirutani vicinity, Higashitonda-mura, Nishi-
muro-gun (ffi^ftlPmi ffl^ilii^ftfe); Wa-
kayama, HONSHU; 33°38'N, 135°26'E; re-
ported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 34:
132
FIELDIANA: ZOOLOGY
XX-26). See Fig. 2A.
Hitotsudai, Okunata-mura, Onyu-gun (3sl!(?P^
=gffl^-7^); Fuktii, HONSHU; 35°23'N,
135°36'E; reported in questionnaire survey
conducted in 1 923 by K. Hasebe (Iwano, 1 974,
p. 39: XXV-9). See Fig. 2A.
Hiyama, Asahi-mura, Adachi-gun ($jM?P)li^
S OJ ); Fukushima, HONSHU; 37°33'N,
140°4rE; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 52:XLI-13). SeeFig. 2A.
Hiyoshi-cho, Funai-gun (fp^?PS"n fflj); Kyoto,
HONSHU; ca. 35°10'N, 135°31'E; collected 9
and 27 Feb. 1980 by JMC staff; JMC, 7 (6
skulls only, 1 skeleton only). Blood samples
collected before 1992 by Nozawa et al. (1991,
p. 414; \996, p. 6). SW38.
Hizaoyama, Shinsaka-son, Jinseki-gun (tt^lP
^^^ BiMiUy, Hiroshima, HONSHU;
34°5rN, 133°16'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 26: XIV-35). See Fig. 2A.
Hodaka (UM); Nagano, HONSHU; ca. 36°17'N,
137°53'E; mtDNA samples collected before
1999 by Y. Kawamoto (1998, p. 54; 2002, p.
60). NE93.
Hodaka Golf Course, Hodaka (^S); Nagano,
HONSHU; 36°18'N, 137°49'E; collected by
PRIKU staff; PRIKU, 1 (skeleton only). NE93.
Hodosan. See Houtosan.
Hokusei-cho, Inabe-gun (M^lPdb^BfT); Mie,
HONSHU; ca. 35°09'N, 136°32'E; mtDNA
samples collected before 2003 by Y. Kawa-
moto (2002, p. 60). SW12.
Hondani National Forest vicinity, Ueno-mura,
Tano-gun ( ^ i?^±i?^;2f^# S #^^);
Gunma, HONSHU; 36°03'N, 138°46'E; re-
ported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 51: XL-1).
See Fig. 2A.
Hongou, Kimitsu-city (:§"/$ TfT ^ 0 ); Chiba,
HONSHU; 35°14'N, 140°00'E; collected 19
Jan. 1999 by PRIKU staff; PRIKU, 1 (skeleton
only). NE120.
Hongou-mura, Ooi-gun (AISIP^^^); Fukui,
HONSHU; 35°28'N, 135°37'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 39: XXV-6). See Fig.
2A.
Hongu-cho, Higashimuro-gun (^#-8?P^'S'
ffll ); Wakayama, HONSHU; ca. 33°50'N,
135°46'E; mtDNA samples collected before
2003 by Y. Kawamoto (2002, p. 60). SW27.
Hongusan, Gakuden-mura, Niwa-gun (i^^lP^
ffl^^^lil); Aichi, HONSHU; 35°19'N,
136°59'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 40: XXVII- 10). See Fig. 2 A.
Hongusan, Ichinomiya-mura, Hoi-gun (^IS^
-t^^^Ol); Aichi, HONSHU; 34°50'N,
137°27'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 40: XXVII- 1). SeeFig. 2 A.
Hongushisan, limuro-son, Asa-gun (^"felPIS^
;|4;zts:$aj); Hiroshima, HONSHU; 34°32'N,
132°25'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 26:XIV-13). SeeFig. 2A.
Honjoyama, Mie-machi, Oono-gun (Ai??PHM
BU ^ J^ Uj ); Oita, KYUSHU; 32°56'N,
131°33'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 16:111-6). See Fig. 2A.
Horane, Seinaiji-mura, Shimoina-gun (T#5P1P
)tl*]S&^)|q|^); Nagano, HONSHU; 35°30'N,
137°48'E; collected 21 Sep. and 19 Oct. 1998
by M. Aimi; PRIKU, 2 (skeletons only). NE72.
Horatani vicinity, Tsurugaoka-mura, Kitaku-
wada-gun (db^BaiPH^ l?^W)l^^ffe); Kyoto,
HONSHU; 35°19'N, 135°33'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 35: XXII- 16). See Fig.
2A.
Hoshio National Forest vicinity, Ozawa-mura,
Kitakanra-gun (db"H'^?PM)H^l.M@W^
fte); Gunma, HONSHU; 36°08'N, 138°39'E;
reported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 51: XL-2).
See Fig. 2A.
Hosoiri-mura, Nei-gun {Wi%M>MX^); Toyama,
HONSHU; ca. 36°32'N, 137°14'E; mtDNA
samples collected before 2003 by Y. Kawa-
moto (2002, p. 60). NE57.
Hotakayama vicinity, Katashina-mura, Tone-gun
(^IJ^lP>^pp^^^li|ffe); Gunma, HONSHU;
36°47'N, 138°50'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 51: XL- 14). See Fig. 2 A.
FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES
133
Hotakayama vicinity. Kavvaba-mura, Tone-gun
{mWi^}\\^^^M\Ui^): Gimma. HONSHU;
36°47'N. 138°50'E; reported in questionnaire
sur\ey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 51: XL- 13). See Fig. 2A.
Hotokedani, Obama-city (d^MTtTi^^); Fiikui.
HONSHU; 35°31'N, 135°43'E; collected Jul.
1994 by PRIKU staff; PRIKU, 1 (postcranials
only). 'SW45.
Hotokeiwa, Hirao-mura, Shimotakai-gun (TM
^ ?P ¥ ^ ^ ^iA ^ ): Nagano, HONSHU;
36°44'N. 138°30'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974. p. 46: XXXI-26). See Fig. 2 A.
Houin, Kisuki-cho, Oohara-gun (AJ^^^/^fflJ
^ ep ); Shimane, HONSHU; 35°13'N,
132°57'E; collected 27 May-12 Dec. 1994 by
PRIKU staff; PRIKU, 3 (skeletons only).
SW59.
Houtakusan. Kamiyoshi-mura, Izushi-gun (ttjS
IP tt H ^ ^ k OJ ); Hyogo. HONSHU;
35°30'N, 134°55'E; reported in questionnaire
sur\ey conducted in 1923 by K. Hasebe
(hvano. 1974. p. 30: XVlII-21). See Fig. 2A.
Houtosan. Nagatoro-machi, Chichibu-gun (^^
^:i:5^fflTi^Uj) (= Hodosan; Nagatoro):
Saitama. HONSHU; 36°04'N, 139°05'E; mon-
key park population, translocated in 1962 from
Gagyusan. Okayama Prefecture (Kanaizuka,
1977, p. 43; cf. Study Group on the Present
Status of Japanese Monkeys, 1977b, p. 23).
Birth season. 1963-1966. reported by Kawai et
al. (1967, p. 39). External measurements taken
before 1997 by Hamada et al. (1996a. pp. 98.
99). NE118.
Houzaw ayama. Kaw akami-mura, Kumano-gun
(^Pipjl|±;^i;RUj); Kyoto, HONSHU;
35°33'N. 134°56'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 35: XXII-6). See Fig. 2A.
Ibuki-cho. Sakata-gun (J^BaiP^R^BO"); Shiga,
HONSHU; ca. 35°23'N. 136°23'E; mtDNA
samples collected before 2003 by Y. Kawa-
moto (2002. p. 60). SW7.
Ibukiyama. Ibuki-mura, Sakata-gun (^ffllP'^RX
^ # R^ OJ ); Shiga, HONSHU; 35°25'N,
136°25'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 38: XXIV- 15). See Fig. 2 A.
Ibukiyama, Samegai-mura, Sakata-gun (^ffl^
M^^^R^OJ)^; Shiga, HONSHU; 35°19'N,
136°23'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano. 1974,
p. 38: XXIV- 18). See Fig. 2A.
Ibukiyama. Suijou-mura. Sakata-gun (^B91P#
Mk^D^Ul); Shiga. HONSHU; 35°23'N,
136°25'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano. 1974,
p. 38: XXIV- 16). See Fig. 2 A.
Ibushikanyama vicinity. Yanadani-mura.
Kamiukena-gun (±>?:iX?P*JP^^-r7^5>t
04^); Ehime, SHIKOKU; 33°31'N, 132°59'E;
reported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano. 1974. p. 21:
IX-2I). SeeFig. 2A.
Ichifusa vicinit}'. Nishimera-son. Ko\ii-gun {%
^^Pe^l^M^TfJMffe); Miyazaki, KYUSHU;
32°I4'N. 131°09'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 15: 11-15). See Fig. 2 A.
Ichiki-cho, Hioki-gun ( B a?PTtT3ifflT); Kago-
shinia. KYUSHU; 31°35'N, 130°21'E; Jomon
subfossils (age, 2.3-12 Ka) reported by K.
Kawanaka (1973, p. 1 18). SW94.
Idodani vicinit\. Shimonokawa-mura,
Ichishi-gun (-^x^T;^JII^#P^ftfe); Mie,
HONSHU; 34°33'N. 136°2rE; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano. 1974. p. 37: XXIII-20). See
Fig. 2A.
Iga-cho. Ayama-gun ( PrI Oj IP # M fiJ ); Mie,
HONSHU; ca. 34°45'N. 136°10'E; collected 23
Oct. 1979-14 Dec. 1983 by PRIKU staff;
PRIKU, 3 (skeletons only). SW20.
Ihama. Minamiizu-cho. Kamo-gun (MMIPM"^
SfflJ^^); Shizuoka, HONSHU; 34°4nsJ,
138°46'E; group fission in 1965 and 1967 re-
ported by Study Group on the Present Status of
Japanese Monkeys (1977b, p. 23). Blood sam-
ples collected before 1992 by Nozawa et al.
(1991. p. 414; 1996.p. 6)..V£/07.
lidesan vicinit>. Toyomi-mura. Higashikan-
bara-gun (^MI^iPMH^tSWOj^); Mi-
gata, HONSHU; 37=41'N, 139=35'E; reported
in questionnaire survey conducted in 1923 by
K. Hasebe (Iwano. 1974. p. 47: XXXII- 18).
See Fig. 2A.
Iketaniyama vicinit>'. Kitagawa-mura, Aki-gun
134
FIELDIANA: ZOOLOGY
($S?PdbJI|^)til#a4fte); Kochi, SHIKOKU;
33°29'N, 134°05'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 22: X-48). See Fig. 2 A.
Ikuno, Doujo-mura, Arima-gun (W/^^PjM^^
^i?); Hyogo, HONSHU; 34°50'N, 135°16'E;
reported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 30:
XVIII-28). See Fig. 2A.
Imaichi-city (^T^TTl:); Tochigi, HONSHU; ca.
36°47'N, 139°39'E; collected 1 Dec. 1984-9
Nov. 1986 by TPM staff; TPM, 8 (skulls only;
excluding 2 skulls not seen, data from speci-
men list). NE43.
Imanishi, Hasumi-mura, Oochi-gun (e^^P^M
W
); Shimane, HONSHU; 34°52'N,
132°37'E; collected 7 Dec. 1996-8 May 1997
by PRIKU staff; KUPR, 6 (skeletons only).
Collected 1 Mar. 2000 by M. Aimi; PRIKU, 1
(skeleton only). SW62.
Inabe-cho, Inabe-gun (M^lPM^fflJ); Mie,
HONSHU; ca. 35°07'N, 136°35'E; mtDNA
samples collected before 2003 by Y. Kawa-
moto (2002, p. 60). SW13.
Inagawa-goryorin, Ookuwa-mura, Nishichi-
kuma-gun (ffim»?PA^^t^i9PJII^3^«);
Nagano, HONSHU; 35°40'N, 137°42'E; re-
ported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 45:
XXXI-5). See Fig. 2A.
Inaodake vicinity, Tashiro-mura, Kimotsuki-gun
( if Jl IP ffl ft W ^ m S ftfe ); Kagoshima,
KYUSHU; 3r07'N, 130°53'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 12: 1-17). See Fig.
2A.
Ine-cho, Yosa-gun {^Wi^^^^); Kyoto,
HONSHU; 35°43'N, 135°17'E; captured alive
5-1 1 Mar. 1980 by JMC staff, died in captivity
6-25 Apr. 1980; JMC, 4 (skulls only). Blood
samples collected before 1992 by Nozawa et al.
(1991, p. 414; \996,^.6).SW48.
Inokamireien, Matsukawa-mura, Kitaazumi-gun
(db^m?P^AMII^^(D#Sg); Nagano,
HONSHU; 36°24'N, 137°49'E; collected 20
Nov. 1998 by PRIKU staff; PRIKU, 1 (skele-
ton only). NE92.
Inouesanrin, Inbi-mura, Minamiamabe-gun (]^
^^P?PHM^#y±Uj^); Oita, KYUSHU;
32°57'N, 131°44'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 16: III- 11). See Fig. 2A.
Inunaki, Kuchimyougata-mura, Gujo-gun (^Ji
^U^^l5¥iii^%); Gifu, HONSHU; 35°46'N,
137°00'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 41 : XXVlII-26). See Fig. 2A.
Inunakisan, Ootsuchi-mura, Sennan-gun (:^^^
A±^;'^lliaj); Osaka, HONSHU; 34°20'N,
135°22'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 33: XIX-6). See Fig. 2A.
Inuyama-city (;^lJL|Tt]); Aichi, HONSHU; ca.
35°23'N, 136°56'N; mtDNA sample collected
before 2003 by Y. Kawamoto (2002, p. 60).
NE6L
Inyou Mountain Range vicinity, Funo-son, Fu-
tami-gun ()5lHlJ^^^^^|iajM-^); Hi-
roshima, HONSHU; 34°52'N, 132°47'E; re-
ported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 27:
XlV-38). See Fig. 2A.
Inyou Mountain Range vicinity, Kimita-son,
Futami-gun ()5(H?P^Ba^ltliajM- w);
Hiroshima, HONSHU; 34°54'N, 132°51'E;
reported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 27:
XIV-37). See Fig. 2A.
Inyou Mountain Range vicinity, Sakugi-son,
Futami-gun {7^^W^^i^^^m\hU-w);
Hiroshima, HONSHU; 34°54'N, 132°44'E;
reported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 27:
XIV-39). See Fig. 2A.
Iriokuyama vicinity, Okouchi-mura, Nishi-
tama-gun (H^ft?P^J^)RlI*l^^^AllLilffe); 7b-
l^o Metropolis, HONSHU; 35°46'N, 139°03'E;
reported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 49:
XXXIV-2). See Fig. 2A.
Irisawai, Ooshika-mura, Shimoina-gun {Y-^M
IP ;^ J^ ^ A )R # ); Nagano, HONSHU;
35°35'N, 138°04'E: collected 7 Dec. 1998 by
PRIKU staff; PRIKU, 1 (fragmented skull
only). NE70.
Iriyamagawaura National Forest, Kuni-mura,
Agatsuma-gun (■
>/N a
nx\h}\mmm
^); Gunma, HONSHU; 36°39'N, 138°40'E;
FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES
135
reported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 51:
XL-10). SeeFig. 2A.
Irokawa-mura, Higashimuro-gun (^^SlPfeJH
^ ); Wakayama, HONSHU; 33°40'N,
135°52'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 34: XX-46). See Fig. 2A.
Isehara-city (^^J^l^J); Kanagawa, HONSHU;
ca. 35°24'N, 139°18'E; mtDNA samples col-
lected before 1998 by Y. Kawamoto (1997, p.
33). NE106.
Ishidosan, Nii-mura, Hikami-gun (7KJL?Plff^
^^P \1\); Hyogo, HONSHU; 35°06'N,
135°02'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 30: XVIII-26). See Fig. 2A.
Ishigagawa, Shimesasu, Nakagawa-mura, Wata-
rai-gun (Jg^lPt JH^jijiJaS ^)R[); Mie,
HONSHU; 34°24'N, 136°32'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 37: XXIII-33). See
Fig. 2A.
Ishizuka, Ichihara-city ( Tfi IP! rfT ^ i^ ); Chiba,
HONSHU; 35°16'N, 140°07'E; collected 1 Sep.
1997-30 Mar. 1999 by PRIKU staff; PRIKU,
4 (skeletons only). NE127.
Isokoen. See Onoaida.
Issou, Kamiyaku-cho, Kumage-gun (^^IPJiM
X^ - My, Kagoshima, YAKUSHIMA;
30°26'N, 130°28'E; collected 25 Feb. 1989 by
PRIKU staff; PRIKU, 1 (skeleton only).
SWIOO.
Issou-Miyanoura. See Yakushima, Census Area
2.
Ita, Heda-mura, Tagata-gun {Wil5WF^¥i^
ffl); Shizuoka, HONSHU; 34°59'N, 138°47'E;
reported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 45:
XXX-17). SeeFig. 2A.
Italy. See Rome.
Itamayama, Saikihon-son, Akaiwa-gun (#^^
feffi^^^W^iUj); Okayama, HONSHU;
34°52'N, 134°07'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 27: XV- 19). See Fig. 2A.
Itotaniyama, Ichiyama-mura, Oochi-gun (e-^^
rtT|l|;^^#Uj); Shimane, HONSHU; 34°57'N,
132°20'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 28:XVI-14). SeeFig. 2A.
Itsukaichi, Akiruno-city ($$^i?rfT£Srfi");
Tokyo Metropolis, HONSHU; ca. 35°44'N,
139°13'E; mtDNA samples collected before
1998 by Y. Kawamoto (1997, p. 33; 2002, p.
6Q).NEU2.
Itsuki-mura, Kuma-gun (^H^S^K^); Ku-
mamoto KYUSHU; 32°24'N, 130°50'E; blood
samples collected before 1992 by Nozawa et al.
(1991, p. 414; \996,^. 6). SW90.
Iwaidani vicinity, Aiga-mura, Kitamuro-gun (db
^«lP*IS^^##ffe); Mie, HONSHU;
34°06'N, 136°13'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 37: XXIII-45). See Fig. 2 A.
Iwaidaniyama, Ibara-mura, Oochi-gun (b^^
#i^^^^^Uj); Shimane, HONSHU;
34°55'N, 132°30'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 28: XVI- 18). See Fig. 2A.
Iwamayama, Ishiyama-mura, Shiga-gun (^M?P
S ill ^^ Pel 111); Shiga, HONSHU; 34°57'N,
135°53'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 38:XXIV-1). SeeFig. 2A.
Iwamiyama National Forest vicinity, Iwami-
sannai-mura, Kawabe-gun (/Rl32.?P^^Hrt
^^.^OlSWWffe); Akita, HONSHU;
39°44'N, 140°23'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 56: XLIV-3). See Fig. 2A.
Iwamiyama vicinity, Toyomatsu-son, Jinseki-gun
( 4$ ^ iP W ^A^ ^ ^ m UJ ffe ); Hiroshima,
HONSHU; 34°48'N, 133°21'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 26: XIV-34). See Fig.
2A.
Iwase National Forest, Yamase-mura, Kita-
akita-gun {it^km^\^M^^MmM^);
Akita, HONSHU; 40°20'N, 140°30'E; reported
in questionnaire survey conducted in 1923 by
K. Hasebe (Iwano, 1974, p. 56: XLIV-7). See
Fig. 2A.
Iwashina-mura, Kamo-gun ( S j^ ^ ^ ^ ^ );
Shizuoka, HONSHU; 34°44'N, 138°48'E; re-
ported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 45:
XXX-23). See Fig. 2A.
136
FIELDIANA: ZOOLOGY
Iwataniyama, Tsuro-mura, Aki-gun ($S^/$S
^^^Uj); Kochi, SHIKOKU; coordinates
unknown; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 22: X-50). Not mapped.
Iwaya, Anabuki-son, Mima-gun (M/^^TvlIl^^
^ M ); Tokushima, SHIKOKU; 34°02'N,
134°10'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 23: XI-25). See Fig. 2A.
Iwayado, Ikeda-son, Yazu-gun (AH^ylfeffl^
^ ^ ^ ); Tottori, HONSHU; 35°19'N,
134°3rE; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 29: XVII-3). See Fig. 2A.
Iwayama, Tarou-mura, Ichishi-gun (— ^IPAfiP
^ ^ ^ Uj ); Mie, HONSHU; 34°32'N,
136°10'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 36:XXIII-18). SeeFig. 2A.
Izu Peninsula. See Hagachizaki.
Jadani vicinity, Yoshinotani-mura, Ishikawa-gun
( ^ JIIIP ^ if ^ ^ !te # ^ ); Ishikawa,
HONSHU; 36°21'N, 136°39'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 40: XXVI-3). See Fig.
2A.
Japan ( 0 ^ ); probably HONSHU; 33°28'-
41°34'E, 130°53'-142°04'E; collected
1823-1829 by P. F. von Siebold (MS.,
1830-1842, p. 1; cf. Temminck, 1842, p. 10;
Holthuis & Sakai, 1970, p. 25); RMNH, 11
(including 4 skins only and 5 skulls only).
Collected 1830-1832 by H. Burger (cf.
Holthuis & Sakai, 1970, p. 37); RMNH, 1
(skeleton only). Specimen received by BM(NH)
from RMNH before 1843, probably collected
1823-1832 by P. F. von Siebold or H. Burger
(cf Gray, 1843, pp. x, 8; 1870, p. 31);
BM(NH), Collected in 1874 by Mr. v.
Hilgendorf; ZMB, 1 (skeleton only). Pur-
chased 1900-1901 by K. A. Haberer (cf
Schweyer, 1909, p. 8); ZSBS, 20 (6 skins only,
14 skulls only). Collected before 1887 by M.
Soller; MNHN, 1. Collected before 1966 by J.
Itani, AIUZ, 2 (skulls only). Not mapped.
Jedo (>IP). See Tokyo.
Jigokudani, Yamanouchi-machi, Shimotakai-gun
(Tit^^POjy rtBTTitki^^) (= Shiga-A);
Nagano, HONSHU; 36°44'N, 138°25'E; birth
season, 1962-1966, reported by Kawai et al.
(1967, p. 38). Provisioning initiated in 1963
(Shidei et al., 1981, p. 16). Field study re-
ported by E. Tokita and S. Hara (1975, p. 24).
Field study conducted 1994-1996 by M.
Iwago and H. Iwago (1999, pp. 154, 160).
NE49.
Jigokudani vicinity, Shoukawa-mura, Oono-gun
(Ai?^SJIIW±tk3i^ffe); Gifu, HONSHU;
35°59'N, 137°10'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 41: XXVIII-38). See Fig. 2 A.
Jinkurousanrin-oku, Hibara-mura, Yama-gun (IP
^iP^J^vWSABPOj^ll); Fukushima,
HONSHU; 37°42'N, 140°02'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 52: XLI-7). See Fig.
2A.
Jinrio, Kamiyama-cho, Myouzai-gun (^ffi^P-f^
lijffll^^) (= Jinryo; Tuino), 150 m elevation;
Tokushima, SHIKOKU; ca. 34°00'N, 134°15'E;
collected 15 and 20 Feb. 1905 by M. P.
Anderson (in Thomas, 1906 ["1905"], p. 336;
cf Kuroda, 1940, p. 272; Napier, 1981, p. 27);
BM(NH), 3. SW70.
Jizoutouge vicinity, Tomikawa-mura, Mi-
namikoma-gun (^^MWmM^ii'^MW^);
Yamanashi, HONSHU; 35°13'N, 138°28'E;
reported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 48:
XXXIII-1). SeeFig. 2A.
Jokohama. See Yokohama.
Jujotouge vicinity, Tomisato-mura, Nishi-
muro-gun (S^«^SM^t + 3tllt#ftfe); Wa-
kayama, HONSHU; 33°45'N, 135°37'E; re-
ported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 33:
XX-21). SeeFig. 2A.
Junigatouge, Nagahama-mura, Minamitsuru-gun
(itl[PmPM^^+-<rllt#); Yamanashi,
HONSHU; 35°29'N, 138°42'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 49: XXXIII-32). See
Fig. 2A.
Kabatani, Kakinoki-mura, Kanoashi-gun {^Sl
^^tK^^^); Shimane, HONSHU; 34°26'N,
131°53'E; reported in questionnaire survey
conducted in 1 923 by K. Hasebe (Iwano, 1 974,
FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES
137
p. 28: XVI-5). See Fig. 2A.
Kabetouge vicinity, Honji-son, Yamagata-gun
(OjmiP^^^oJ^llfF^); Hiroshima,
HONSHU; 34°39'N, 132°32'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 26: XIV- 10). See Fig.
2A.
Kabetouge vicinity, Minamikata-son, Yama-
gata-gun {\h%W^l5^'^U^^)\ Hi-
roshima, HONSHU; 34°39'N, 132°34'E; re-
ported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 26:
XIV-U). SeeFig. 2A.
Kabutoyama, Kabuto-mura, Suzuka-gun (Ip^?P
iPA^iPAUj); Mie, HONSHU; 34°52'N,
136°2rE; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 36:XXIII-11). SeeFig. 2A.
Kado-mura, Ooi-gun (AtS^iP4^); Fukui,
HONSHU; 35°29'N, 135°40'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 39: XXV-8). See Fig.
2A.
Kagamisakadani, Oono-mura, Kitakuwada-gun
(db#BaiPAifW^^#); Kyoto, HONSHU;
35°16'N, 135°3rE; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 35: XXII-14). See Fig. 2A.
Kaisho, Ootaki, Asahi-machi, Nishimura-
yama-gun (ffiWUj^^ S BJ A)i^Pif); Ya-
magata, HONSHU; 38°16'N, 140°07'E; col-
lected 15 Oct. 1997 by PRIKU staff; PRIKU, 2
(skeletons only). NE30.
Kakeban vicinity, Sawatani-son, Naka-gun {M%.
fP)i#WShMffe); Tokushima, SHIKOKU;
33°5rN, 134°14'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 23: XI-14). See Fig. 2A.
Kakisore, Nagiso-machi, Kiso-gun (^h?P1^^
^fflj^^); Nagano, HONSHU; 35°39' N,
137°37'E; collected 30 Aug. and 3 Sep. 1998
by M. Aimi; PRIKU, 2 (skeletons only). NE75.
Kakuma National Forest, Osa-mura, Chiisa-
gata-gun ('\^W^WB:^^f^^^^y, Nagano,
HONSHU; 36°26'N, 138°22'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 46: XXXI-32). See
Fig. 2A.
Kakumahachi, Matsukura-mura, Shimonii-
kawa-gun {T^JWW'^ii'^^MBUy, Toyama,
HONSHU; 36°45'N, 137°27'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 42: XXIX-7). See Fig.
2A.
Kamae-machi, Minamiamabe-gun (^^nP^M
>IfflT); Oita, KYUSHU; 32°48'N, 131°56'E;
blood samples collected before 1992 by No-
zawa et al. ( 1 99 1 , p. 4 1 4; 1 996, p. 6). SW82.
Kamagatani, Kamiijira-mura, Yamagata-gun (ill
m?P±#gm^ll<r#); Gifu, HONSHU;
35°32'N, 136°44'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 41 : XXVIII- 12). See Fig. 2A.
Kamanashiyama, Ochiai-mura, Suwa-gun (MIS
^P^-^^H^Lij); Nagano, HONSHU;
35°52'N, 138°17'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 46: XXXI-40). See Fig. 2A.
Kametaniyama vicinity, Michikawa-mura,
Mino-gun {^1mWM.}\\¥i%.'^\h^); Shimane,
HONSHU; 34°38'N, 132°06'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 28: XVI- 13). See Fig.
2A.
Kameyama-city (MUlTtT); Mie, HONSHU; ca.
34°51'N, 136°27'E; mtDNA samples collected
before 2003 by Y. Kawamoto (2002, p. 60).
SWI8.
Kamidaira, Takamori ("F^S[51PS^fflT±¥);
Nagano, HONSHU; 35°33'N, 137°50'E; col-
lected 12 Dec. 1997-18 Aug. 1998 by M. Aimi;
PRIKU, 30 (skeletons only). NE76.
Kamigamo, Kyoto-city (i^tPrfTJiM^); Kyoto,
HONSHU; 35°03'N, 135°45'E; collected 24
Jul. 1998 by PRIKU staff; PRIKU, 1 (skeleton
only). SW37.
Kamiichi-machi, Nakaniikawa-gun (4'WfJII^-t
TtJ fflj ); Toyama, HONSHU; ca. 36°42'N,
137°22'E; mtDNA samples collected before
2003 by Y. Kawamoto (2002, p. 60). NE55.
Kamikaifu-mura, Iwafune-gun (^^plP-h^i^
^); Niigata, HONSHU; 38°18'N, 139°28'E;
reported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 47:
XXXII-22). See Fig. 2A.
Kamikitayama-mura, Yoshino-gun (n^i??P_bdb
OjW); Nara, HONSHU; 34°09'N, 136°01'E;
reported in questionnaire survey conducted in
138
FIELDIANA: ZOOLOGY
1923 by K. Hasebe (Iwano, 1974, p. 35:
XXI-6). See Fig. 2A.
Kamikochi, Azumi-mura, Minamiazumi-gun (^
$»?P^*W±I^±tk); Nagano, HONSHU;
ca. 36°15'N, 137°38'E; mtDNA samples col-
lected before 1998 by Y. Kawamoto (1997, p.
33). NE87.
Kamikumatani, Kumatani-son, Atetsu-gun (PrI^
?P^#^±^^); Okayama, HONSHU;
35°0rN, 133°3rE; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 27: XV- 10). See Fig. 2A.
Kamimachi, Kami-mura, Shimoina-gun (T^M
?P±:^±fflT); Nagano, HONSHU; 35°23'N,
137°58'E; collected 17 Dec. 1997 by M. Aimi;
PRIKU, 1 (skeleton only). NE69.
Kamimobiki vicinity, Mobiki-mura, Kimo-
tsuki-gun (FMlPW?l^±^?lfte); Kago-
shima, KYUSHU; 31 °29'N, 130°54'E; reported
in questionnaire survey conducted in 1923 by
K. Hasebe (Iwano, 1974, p. 12: 1-13). See Fig.
2A.
Kaminada, Nakatsugawa-gumi, Sumoto-city,
Awajishima ()iili^ A}J+I^TtJ4^}$JI||i±il);
Hyogo. AWAJISHIMA; 34°14'N, 134°52' E;
field study reported by M. Nakamichi (1989, p.
738; cf. Nakamichi et al., 1997, p. 227). SW69.
Kaminaka-cho, Onyu-gun (islij^P-h't'fflT); Fu-
kui, HONSHU; 35°27'N, 135°51'E; collected
Oct.-Dec. 1994 by PRIKU staff; PRIKU, 3
(skeletons only). SW44.
Kaminoyama-city ( Ji |1| Tti ); Yamagata,
HONSHU; ca. 38°14'N, 140°17'E; mtDNA
samples collected before 2003 by Y. Kawa-
moto (2002, p. 60). NE29.
Kamioota-mura, Higashimuro-gun (^#-ftlP_h
;<!; Ba W ); Wakayama, HONSHU; 33°37'N,
135°53'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 34: XX-44). See Fig. 2A. ^
Kamisanji-mura, Hidaka-gun ( S i^?P_h|llS&^);
Wakayama, HONSHU; 33°55'N, 135°31'E;
reported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 33:
XX-12). SeeFig. 2A.
Kamisawayama, Kisawa-mura, Shimoina-gun
( T ^ SP IP tK )R ^ ± >R Oj ); Nagano,
HONSHU; 35°21'N, 137°58'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 47: XXXI-48). See
Fig. 2A.
Kamiseinaiji, Seinaiji-mura, Shimoina-gun (T#
MWm\k^^ ±M\H^); Nagano, HONSHU;
35°30'N, 137°4rE; collected 28 Oct. 1998 by
M. Aimi; PRIKU, 1 (skeleton only). Collected
25 Nov. 1998 by PRIKU staff; PRIKU, 1
(fragmented skull only). NE73.
Kamitakarazawa, Higashisawa-mura, Minami-
murayama-gun (]^^ Uj ?P^}R:14±S)H);
Yamagata, HONSHU; 38°12'N, 140°27'E; re-
ported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 55:
XLII-13). SeeFig. 2A.
Kamitakashiri, Nanukaichi-mura, Kanoashi-gun
H^^m^Z^l^^hM^y, Shimane,
HONSHU; 34°23'N, 131°55'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 28: XVI-4). See Fig.
2A.
Kamiyaku-cho [1], Kumage-gun (^^lP_hM^
fflj); Kagoshima, YAKUSHIMA; ca. 30°25'N,
I30°35'E; blood samples collected before 1992
by Nozawa et al. (1991, p. 414; 1996, p. 7).
SWIOO.
Kamiyaku-cho [2], Kumage-gun (rI^^PJiM^
HTT); Kagoshima, YAKUSHIMA; 30°16'N,
130°29'E; collected 26 Nov. 1977 by PRIKU
staff; PRIKU, 1 (skeleton only). Collected 9
Jun. 1989 by S. Azuma; PRIKU, 14 (skeletons
only). SWIOI.
Kamou, Kimitsu-city ( ^ >$ r|T H ^ ); Chiba,
HONSHU; 35°13'N, 140°07'E; collected 30
Jul. 1996-30 Mar. 1999 by PRIKU staff;
PRIKU, 7 (skeletons only, including 1 with
fragmented skull). NE126.
Kanakidoyama, Kamitakara-mura, Yoshiki-gun
(^^?P±S^^7KFlij); Gifu, HONSHU;
36°20'N, 137°25'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 41: XXVIII-36). See Fig. 2A.
Kanasumi National Forest, Kijo-son, Koyu-gun
(Jft^iPTK^^J^J^aW^); Miyazaki,
KYUSHU; 32°14'N, 131°24'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 15: 11-18). See Fig.
2A.
Kanatokoyama, Aihashi-mura, Izushi-gun (tij^
IP-^M^^iiUj); Hyogo, HONSHU;
FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES
139
35°24'N, 134°56'E; reported in questionnaire
sur\ey conducted in 1923 by K. Hasebe
(hvano, 1974, p. 30: XVIII- 19). See Fig. 2 A.
Kanatokoyama, Itoi-mura, Yabu-gun (M5^1P^
^^^iiUl); Hyogo. HONSHU; 35°2rN,
134°53'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 30:XVIII-16). SeeFig. 2A.
Kanaya-cho. Arida-gun (WBaiP^MfflJ): Wa-
kayama. HONSHU; ca. 34°04'N, 135°15'E;
mtDNA samples collected before 2003 by Y.
Kawamoto (2002. p. 60). SW31.
Kanayama. Satsuma-cho. Satsuma-gun (SS?P
SSfflJ^Uj): Kagoshima. KYUSHU; 31°55'N.
130°36'E; collected 24 Jan. 2000 by M. Aimi;
PRIKU. 5 (skulls only). SW92.
Kanazavvayama \icinity. Miyagase-mura.
Aiko-gun ( g ¥ ?P ^ V ^ ;^ ±^R Uj ^ );
Kanagaua, HONSHU; 35°31'N, 139°14'E;
reported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano. 1974. p. 50:
XXXV-7). See Fig. 2A.
Kanba. Katsuyama-cho. Maniwa-gun (XfilPM
Lijffllttfi); Okayama. HONSHU; 35°05'N.
133°42'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano. 1974.
p. 27:XV-13). SeeFig. 2A.
Kanbanotaki. Katsuyama-cho. Maniwa-gun (Ml
JS?PBlijfflIttJS(7)>t); Okayama. HONSHU;
35°07'N. 133°4rE; provisoned group; birth
season. 1962. reported by Kawai et al. (1967.
pp. 37. 38). SW55.
Kandayama. Hongou-cho. Kitashitara-gun (dbix
^A^iPfflTttBaOJ); AichL FIONSHU;
35°05'N. 137°42'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano. 1974. p. 40: XXVII-4). See Fig. 2 A.
Kanegaryumorisankei. Nakayama-mura. Aki-gun
( $^^' ?P ^ 111 ;^ M <r ^ ^ UJ .^ ); Ko^chL
SHIKOKU; 33°30'N, 134°02'E; reported in
questionnaire sur\ey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 22: X-49). See Fig.
2A.
Kanegatake \icinity. Kiyokawa-mura. Naka-
koma-gun (4^ ^SlPTf JH^^ ^ S^); Ya-
manashi, HONSHU; coordinates unknown;
reported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 48:
XXXIII-13). Not mapped.
Kanehirayama, Hirakawa-son, Kawakami-gun
(JI|±?P¥JI|^^¥aj); Okayama, HONSHU;
34°49'N. 133°25'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano. 1974. p. 27: XV-3). See Fig. 2A.
Kaneyamadani. Kuragi-mura. Kanoashi-gun (^
S-IPM^W^OJ^); Shimane, HONSHU;
34°22'N, 131°59'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 28: XVI- 1). See Fig. 2 A.
Kanita-machi. Higashitsugaru-gun (!^/$fe?PS
Ba fflj ); Aomon. HONSHU; ca. 41°02'N,
140°39'E; mtDNA samples collected before
1998 by Y. Kawamoto (1997. p. 33: 2002. p.
60). NE6.
Kanmuriyama. Mizusawa-mura. Mie-gun (HM
IP7K)R:ftEUj); Mie. HONSHU;^ 34°57'N,
136°28'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974.
p. 36: XXIII-6). See Fig. 2A.
Kanmuriyama. Takahara-mura. Oochi-gun (e^
W>Md^MlU): Shimane, HONSHU; 34°55'N,
132°34'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 28: XVI- 19). SeeFig. 2A.
Kannon. See Kannonzaki.
Kannonyama. Yotsu-mura. Takaoka-gun (Si^
IP ^')$ ^ ^ # OJ ); KochL SHIKOKU;
33°irN, 133°13'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974. p. 22: X-16). See Fig. 2A.
Kannonzaki. Kamiyaku-cho. Kumage-gun {M.^
lP±M^ffltia^*f); ^Kagoshima.
YAKUSHIMA; ca. 30°23'N, 130°23'E; ob-
served Oct. 1977 by T. Maruhashi (1982, p.
318; cf. Takasaki & Masui. 1984, p. 311).
SWIOI.
Kanotoyama vicinity. Hinohara-mura, Nishi-
tama-gun (S^0iP^JI^#P Ujffe); Tokyo
Metropolis. HONSHU; 35°43'N, 139°07'E;
reported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 49:
XXXIV-1). SeeFig. 2A.
Kanoya-city (^MTfT); Kagoshima. KYUSHU;
ca. 31°22'N, 130°46'E; mtDNA samples col-
lected before 2003 by Y. Kawamoto (2002, p.
60). SW95.
Karamatsu, Miyoshi-son, Atetsu-gun (Po[^^||
^^0^i); Okayama, HONSHU; 34°58'N,
140
FIELDIANA: ZOOLOGY
133°3rE; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 27:XV-11). SeeFig. 2A.
Karasugawa, Nishihodaka-mura, Minamia-
zumi-gun (]^$ft?PffiM>S^,^JI|); Nagano,
HONSHU; 36°19'N, 137°51'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 46: XXXI- 15). See
Fig. 2A.
Karasugawa National Forest, Karasugawa-mura,
Minamiazumi-gun ( j% ^ S IP B J i I W .i J 1 1 S
#^); Nagano, HONSHU; 36°16'N, 137°52'E;
reported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 46:
XXXI-14). SeeFig. 2A.
Karekidani, Higashiyashiki, Shikiya-mura, Hi-
gashimuro-gun (^^W:^WiE.^%M.Wi^^
^ ); Wakayama, HONSHU; 33°50'N,
135°49'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 34:XX-51). SeeFig. 2A.
Karioyama vicinity, Yahata-son, Yamagata-gun
( Ol m ?P A 1i ^ XIJ M UJ ffe ); Hiroshima,
HONSHU; 34°41'N, 132°11'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 26: XIV-6). See Fig.
2A.
Karitayama, Takayama-mura, Kamitakai-gun (_h
m^W>m\^¥iM^\h); Nagano, HONSHU;
36°41'N, 138°20'E; collected 1 Nov. 1998 by
PRIKU staff; PRIKU, 1 (skeleton only). NE49.
Karuizawa-machi, Kitasaku-gun (db'fe^^S^
>RlflT); Nagano, HONSHU; ca. 36°21'N,
138°38'E; mtDNA samples collected before
2003 by Y. Kawamoto (2002, p. 60). NE45.
Kasagi, Kashiwazaki-mura, Watarai-gun (S^
^^fillit^^TK); Mie, HONSHU; 34°18'N,
136°23'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 37: XXIII-30). See Fig. 2A.
Kashiji vicinity, Ichiu-son, Mima-gun (H/^IP —
^ t^ @ itk fife ); Tokushima, SHIKOKU;
33°56'N, 134°05'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 23: XI-23). See Fig. 2A.
Kashima (Eft); Ehime, KASHIMA; 32°57'N,
132°27'E; onset of provisioning in 1955 re-
ported by S. Yoshihiro (1983, p. 41; cf Edito-
rial Committee of Nihonzaru, 1977, p. 112).
Blood samples collected before 1992 by No-
zawa et al. (1991, p. 414; 1996, p. 6). SW81.
Kashima, Nishisotoumi-mura, Minamiuwa-gun
(l^^^^lPS^^^Jift); Ehime, KASHIMA
32°57'N, 132°27'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 18: IX- 1). See Fig. 2A.
Kashima-machi, Soma-gun ( ^1 M IP E ft BJ );
Fukushima, HONSHU; ca. 37°42'N, 140°58'E;
mtDNA samples collected before 2003 by Y.
Kawamoto (2002, p. 60). NE59.
Kashio, Ooshika-mura, Shimoina-gun (T#i§P?P
AMWMi^); Nagano, HONSHU; 35°36'N,
138°03'E; collected 18 Dec. 1997 by M. Aimi;
PRIKU, 1 (skeleton only). Collected 14 Mar.
1998 by PRIKU staff; PRIKU, 1 (skeleton
only). NE70.
Kashiwabarayama foothills, Kaminada-mura,
Tsuna-gun (^=glP±yllWttJ^ajM); Hyogo,
AWAJISHIMA; 34°16'N, 134°53'E; reported
in questionnaire survey conducted in 1923 by
K. Hasebe (Iwano, 1974, p. 30: XVIII-2). See
Fig. 2A.
Kashiwajima, Okuuchi-mura, Hata-gun ("l^^^
Urt^ttft); Kochi, SHIKOKU; 32°47'N,
132°40'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 21:X-1). SeeFig. 2A.
Kashiyama vicinity, Funatsuki-mura, Hidaka-gun
( B S IP ^q' * ^ a Ol ftfe ); Wakayama,
HONSHU; 33°57'N, 135°17'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 33: XX-7). See Fig.
2A.
Kasshizan, Nishigou-mura, Nishishirakawa-gun
O a )Rl IP a ^ ;^^f ¥ ^ UJ ); Fukushima,
HONSHU; 37°irN, 140°09'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 52: XLl-5). See Fig.
2A.
Kasuga-cho, Hikami-gun ( tR ± IP # 0 fflj )
Hyogo, HONSHU; ca. 35°09'N, 135°10'E
collected 23 and 27 Feb. 1980 by JMC staff;
JMC, 7 (5 skulls only, 2 skeletons only). Blood
samples collected before 1992 by Nozawa et al
(1991, p. 414; 1996, p. 6). SW40.
Kasugadaira, lijima-machi, Kamiina-gun (_h'^
3P?PI5ftfflT#B¥); Nagano, HONSHU;
35°42'N, 137°54'E; collected 30 Oct. and 2
FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES
141
Dec. 1997 by M. Aimi; PRIKU, 4 (skeletons
only, including 2 with fragmented skulls).
Collected 12 Nov. and 2 Dec. 1997 by PRIKU
staff; PRIKU, 2 (skeletons only, fragmented
skulls). NE77.
Kasuge-mura, Yamamoto-gun (|1|^^^^^);
Akita, HONSHU; 40°17'N, 140°18'E; reported
in questionnaire survey conducted in 1923 by
K. Hasebe (Iwano, 1974, p. 56: XLlV-1 1). See
Fig. 2A.
Kataiwa National forest, Kamigou-mura, Kami-
hei-gun (±r^#?P±^^>^^aWW); Iwate,
HONSHU; 39°15'N, 141°36'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Mito, 1989, p. 25). See Fig. 2A.
Katamukiyama, Hakusan-mura, Oono-gun (Ai?
^PfiOj^^Oj); Oita, KYUSHU; 32°49'N,
131°30'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 16:111-4). See Fig. 2A.
Kataoka-mura, Ika-gun ('^^^jnf^^^); Shiga,
HONSHU; 35°33'N, 136°12'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 38: XXIV- 11). See
Fig. 2A.
Katsuojisan, Toyokawa-mura, Mishima-gun (H
IIPWJH^SM^UJ); Osaka, HONSHU;
34°50'N, 135°33'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 33: XIX-3). See Fig. 2A.
Katsurashima, Nakagawa-mura, Kamiina-gun
(±#i9P?P4'JII^Sft); Nagano, HONSHU;
35°36'N, 137°56'E; collected 7 Jul. 1998 by
PRIKU staff; PRIKU, 1 (skeleton only). NE71.
Katsuyama-cho, Maniwa-gun (M^lPMlilfflJ);
Okayama, HONSHU; 35°05'N, 133°38'E;
blood samples collected before 1992 by No-
zawa et al. (1991, p. 414; 1996, p. 6). Field
study reported by Itoigawa et al. (1992, p. 58).
External measurements taken before 1997 by
Hamada et al. (1996a, pp. 98, 99). SW54.
Kawabe-machi, Kawabe-gun (yRlaS^PyRjiSfflJ);
Akita, HONSHU; ca. 39°38'N, 140°13'E;
mtDNA extracted from hunting trophies before
2003 by T. Agatsuma and M. Ishigami (2002,
p. 79). NE14.
Kawachi vicinity, Mugi-cho, Kaifu-gun (^pP?P
# tt fflj )Rr rt flk ); Tokushima, SHIKOKU;
33°40'N, 134°26'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 23: XI-9). See Fig. 2 A.
Kawachiyama, Kawachi-mura, Nakakanbara-gun
(4^aj^^JI||*l^JllrtUj); Niigata, HONSHU;
37°4rN, 139°14'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 47: XXXIl-15). See Fig. 2A.
Kawaguchi-ko ()RrP)^); Yamanashi, HONSHU;
35°31'N, 138°45'E; two groups, enclosed and
provisioned before 1978, reported by Study
Group on the Present Status of Japanese Mon-
keys (1977b, p. 23). Translocation from
Kamae and Shodoshima reported by T. Shidei
etal. (1981,p. \1).NE97.
Kawajionsen, Fujiwara-machi, Shioya-gun {^^
?P^J^fflIJI|>^;U;7^); Tochigi, HONSHU;
36°53'N, 139°42'E; collected 8 Dec. 1986 by
TPM staff; TPM, 1 (not seen, data from
specimen list). NE41.
Kawakami-mura, Yoshino-gun CRiJ^PJH-h^);
Nara, HONSHU; 34°18'N, 136°00'E; reported
in questionnaire survey conducted in 1923 by
K. Hasebe (Iwano, 1974, p. 35: XXI-7). See
Fig. 2A.
Kawamatasanrin, Mazuma-mura, Hidaka-gun
( B ift ?P M S W Jll 3104 ^ ); Wakayama,
HONSHU; 33°51'N, 135°19'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 33: XX- 10). See Fig.
2A.
Kawanaka-mura, Hidaka-gun (0S^JII4'W);
Wakayama, HONSHU; 33°56'N, 135°20'E;
reported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 33: XX-8).
See Fig. 2A.
Kawanedake National Forest vicinity, Kawa-
saki-mura, Shibata-gun (^Ba?PJI|lli^WJII a"^
aWWffe); Miyagi, HONSHU; 38°11'N,
140°38'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 55: XLlII-5). See Fig. 2A.
Kawanishi, Matsukawa-mura, Kitaazumi-gun (db
^•?P;^iJIIWJI|ffi); Nagano, HONSHU;
36°26'N, 137°50'E; collected 22 Feb. 1998 by
M. Aimi; PRIKU, 2 (skeletons only). NE92.
Kawara, Kamiyaku-cho, Kumage-gun (^^^-h
M:^fflTJI|J^); Kagoshima, YAKUSHIMA; ca.
30°21'N, 130°23'E; observed Jul. 1975 by T.
Maruhashi (1982, p. 318; cf. Takasaki & Ma-
142
FIELDIANA: ZOOLOGY
sui, 1984, p. 311). 5^7 r?/.
Kawaradake (##S); Fukuoka, KYUSHU;
33°40'N, 130°51'E; blood samples collected
before 1992 by Nozawa et al. (1991, p. 414;
1996,p. 6).5Pf<^^.
Kawaradake, Kawara-machi, Tagawa-gun (B9JII
iP##fflT##|5); Fukuoka, KYUSHU;
33°28'N, 130°44'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 17: VII-2). See Fig. 2 A.
Kawaratai vicinity, Nishimeya-mura, Na-
katsugaru-gun (4^)*felPffii Jl^Jlli^¥ffe);
Aomori, HONSHU; 40°3rN, 140°16'E; re-
ported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 57:
XLVI-7). See Fig. 2A.
Kawarayama, Tsutasawa-mura, Shisou-gun (5^
^ 15 M )R W )Rl Jl Lij ); Hyogo, HONSHU;
35°02'N, 134°3rE; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 30: XVIII-6). See Fig. 2A.
Kawasuso, Seinaiji-mura, Shimoina-gun {~^^M
^m\H^^]\\^); Nagano, HONSHU;
35°28'N, 137°42'E; collected 19 Oct. 1998 by
PRIKU staff; PRIKU, 1 (skeleton only). NE73.
Kawatayama vicinity, Miyama-son, Oe-gun (^
ffl^HLl4;f4Jllfflajffe); Tokushima,
SHIKOKU; 34°00'N, 134°14'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 23: XI-27). See Fig.
2A.
Kawaurayama, Itadori-mura, Mugi-gun (^'ffi^P
^IX^^JIhHOj); Gifu, HONSHU; 35°44'N,
136°48'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 41:XXV1II-16). SeeFig. 2A.
Kayano vicinity, Oshiro-mura, Mikata-gun (H^
^ dMt ^ ^ i? -te ); Hyogo, HONSHU;
35°25'N, 134°30'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 30: XVllI-10). See Fig. 2A.
Kayano-Nanamagari, Kisarazu-city (^M/$rfi^
i? -b te ); Chiba, HONSHU; 35°20'N,
140°05'E; collected 6 Nov. 1996-12 Jan. 1999
by PRIKU staff; PRIKU, 3 (skeletons only, in-
cluding 1 with fragmented skull). NE128.
Kensozan, Kitaura-mura, Shozu-gun (d'^S.lPdb
)i ^^t lll^ llil 111 ); Kagawa, SHODOSHIMA;
34°32'N, 134°16'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 24: XII-7). See Fig. 2A.
Kensozan, Kusakabe-mura, Shozu-gun (d'^SlP
^Mt^lllJ^lliflOj); Kagawa, SHODOSHIMA;
34°30'N, 134°19'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 24: XII-9). See Fig. 2A.
Kensozan, Oobe-mura, Shozu-gun (<J'^a.^AnP
^ i!^ Ilifl Uj ); Kagawa, SHODOSHIMA;
34°32'N, 134°19'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 24: XII-8). See Fig. 2A.
Kensozan, Oonude-mura, Shozu-gun (-'J'^S.^A
^ ^ ll^ lli§ UJ ); Kagawa, SHODOSHIMA;
34°30'N, 134°15'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 24: XII-6). See Fig. 2 A.
Kensozan, Yasuda-mura, Shozu-gun {'\'^W^^
Ba ;^>)- ll!^ Ilil Ol ); Kagawa, SHODOSHIMA;
34°29'N, 134°21'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 24: XII- 10). See Fig. 2A.
Kidani, Natasho-mura, Onyu-gun {^%M>^ B9
Ji ^ ^ # ); Fukui, HONSHU; 35°23'N,
135°43'E; collected Jul. 1994 by PRIKU staff;
PRIKU, 2 (skeletons only, including 1 with
fragmented skull). SW42.
Kidooku, Ebara-son, Mima-gun (H/^IP/IJ^W
?KFII); Tokushima, SHIKOKU; 34°08'N,
134°irE; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 23:XI-31). SeeFig. 2A.
Kigiura vicinity, Mikita-cho, Kaifu-gun (^nPlP
Hllij^BafflT7KlliJ^)ifte); Tokushima, SHIKOKU;
33°47'N, 134°36'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 23: XI-6). See Fig. 2 A.
Kimitsu-city (^^Tfi); Chiba, HONSHU; ca.
35°19'N, 139°53'E; provisioning 1969-1976
reported by Study Group on the Present Status
of Japanese Monkeys (1977b, p. 26). MtDNA
samples collected before 1999 by Y. Kawa-
moto (1998, p. 54). ^£7/ 9.
Kinbaratani, Toyama-mura, Motosu-gun (^^?P
^KUJ^^J^^); Gifu, HONSHU; 35°34'N,
136°39'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 41:XXV111-11). SeeFig. 2A.
Kinkazan (^llOj); Miyagi, KINKAZAN (=
FOODEN AND AIMI: SYSTEMATIC REVIEW OE JAPANESE MACAQUES
143
KINKASAN); 38°16'-38°19'N, 141°33'-
141°36'E; birth season, 1966, reported by
Kawai et al. (1967, p. 39; cf. Study Group on
the Present Status of Japanese Monkeys, 1977a,
p. 10). Collected 11 Aug. 1982 by K. Izawa;
MUE, 1 (skull only). MtDNA samples col-
lected before 1998 by Y. Kawamoto (1997, p.
33; 2002, p. 60). Not mapped.
Kinkazan vicinity, Ayukavva-mura, Oshika-gun
( tt E IP l£ Jll W ^ H OJ fte ); Mvfl^/,
KINKAZAN; 38°18'N, 141°33'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 55: XLIlI-12). See
Fig. 2A.
Kinkazan, Loc. 1 ( ^ H ill ); Miyagi.
KINKAZAN; 38°17.5'N, 141°35.5'E; collected
3 Dec. 1982 by K. Izawa; MUE, 1 (skull only).
NE22.
Kinkazan, Loc. 2 ( :^ H |JL| ); Miyagi.
KINKAZAN; 38°18.25'N, 141°35'E; collected
18 Jul. 1983 by K. Izawa; MUE, 1 (skull only).
NE22.
Kinkazan, Loc. 6 ( :^ ¥ ill ); Miyagi,
KINKAZAN; 38°17.5'N, 141°34'E; collected
24 Feb. 1984 by K. Izawa; MUE, 1 (skull
only). NE22.
Kinkazan, Loc. 10 ( ^ H |JL| ); Miyagi.
KINKAZAN; 38°17.25'N, 141°34'E; collected
31 Mar. 1984 by K. Izawa; MUE, 1 (skull
only). NE22.
Kinkazan, Loc. 11 ( :^ H ill ); Miyagi.
KINKAZAN; 38°18.25'N, 141°35'E; collected
7 Apr. 1984 by K. Izawa; MUE, 1 (skull only).
NE22.
Kinkazan, Loc. 12 ( :^ ¥ ill ); Miyagi.
KINKAZAN; 38°17.25'N, 141°34'E; collected
14 Apr. 1984 by K. Izawa; MUE, 1 (skull
only). NE22.
Kinkazan, Loc. 13 ( ^ H ill ); Miyagi.
KINKAZAN; 38°18'N, 141°34'E; collected 23
Apr. 1984 by K. Izawa; MUE, 1 (skull only).
NE22.
Kinkazan, Loc. 14 ( :^ ¥ ill ); Miyagi,
KINKAZAN; 38°18'N, 141°34'E; collected 29
Apr. 1984 by K. Izawa; MUE, 1 (skull only).
NE22.
Kinkazan, Loc. 15 ( :^ ^ ill ); Miyagi.
KINKAZAN; 38°17'N, 141°35'E; collected 17
May 1984 by K. Izawa; MUE, 1 (skull only).
NE22.
Kinkazan, Loc. 20 ( :#: H ill ); Miyagi,
KINKAZAN; 38°17.5'N, 141°35'E; collected
20 Jan. 1985 by K. Izawa; MUE, 1 (skull only).
NE22.
Kinkazan, Loc. 24 ( ^ H |lj ); Miyagi,
KINKAZAN; 38°16.5'N, 141°35.5'E; collected
15 May 1985 by K. Izawa; MUE, 1 (skull
only). NE22.
Kinkazan, Loc. 28 ( ^ H ill ); Miyagi,
KINKAZAN; 38°17.25'N, 141°34'E; collected
12 Apr. 1986 by K. Izawa; MUE, 1 (skull
only). NE22.
Kinkazan, Loc. 33 ( :^ ? ill ); Miyagi,
KINKAZAN; 38°17'N, 141°35.5'E; collected 3
Oct. 1988 by K. Izawa; MUE, I (skull only).
NE22.
Kinkazan, Loc. Y6 ( :^ ? ill ); Miyagi,
KINKAZAN; 38°17.25'N, 141°35'E; collected
26 May 1984 by K. Izawa; MUE, 1 (skull
only). hE22.
Kinpusan vicinity, Miyamoto-mura, Naka-
koma-gun {^^W.^'M^^±^\^^); Ya-
manashi, HONSHU; 35°51'N, 138°37'E; re-
ported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 48:
XXXIII-12). SeeFig. 2A.
Kinsenji. See Kyogatake.
Kintokiyama, Ashigara-mura, Sunto-gun (f^M
?P £ ffi ^ ^ B#'UJ ); Shizuoka, HONSHU;
35°19'N, 139°00'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 45: XXX- 15). See Fig. 2A.
Kintokiyama vicinity, Sengokuhara-mura, Ashi-
garashimo-gun (£ffiT#fllj^I^W^Btlljflfe);
Kanagawa, HONSHU; 35°17'N, 139°0rE;
reported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 49:
XXXV-4). See Fig. 2A.
Kirakuyama vicinity, Kiyomizu-mura,
Agawa-gun (#JlliP^7K;^^Maj^); Kochi,
SHIKOKU; 33°44N, 133°22'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 22: X-26). See Fig.
2A.
Kirigatana, Touni-mura, Kesen-gun (^"(llj^^
:^^^ fl^ -Jr il ); luate, HONSHU; 39°12'N,
141°5rE; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
144
FIELDIANA: ZOOLOGY
p. 56: XLV-6). See Fig. 2A.
Kirigataniyama vicinity, Kawagoe-mura,
Oochi-gun {G^m}\m^m'T'^aii\k): Shi-
mane, HONSHU; 34°57'N, 132°28'E; reported
in questionnaire survey conducted in 1923 by
K. Hasebe (Iwano, 1974, p. 28: XVI- 17). See
Fig. 2A.
Kirishimasanrin, Higashisonoyama-mura, Aira-
gun (Itp^^P^ftOJ^^tSftUj^); Kagoshima,
KYUSHU; 31°50'N, 130°52'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 12: I- 10). See Fig.
2A.
Kirishimayama, Nishidake-son, Kitamoro-
kata-gun (itMW^^^^^MMiU); Miyazaki,
KYUSHU; 31°52'N, 130°58'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 15: II-8). See Fig. 2 A.
Kirishimayama vicinity, Takahara-son, Ni-
shimorokata-gun {'^WW.^mW-^^BMiUifky,
Miyazaki, KYUSHU; 3r56'N, 131°04'E; re-
ported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 15: 11-12).
See Fig. 2A.
Kishida, Hatta-mura, Mikata-gun (M^^fPABB
^^f # ffl ); Hyogo, HONSHU; 35°31'N,
134°26'E; reported in questionnaire survey
conducted in 1 923 by K. Hasebe (Iwano, 1 974,
p. 30:XVIII-11). SeeFig. 2A.
Kiso ( tK ^ ); Nagano, HONSHU; 35°45'N,
137°43'E; field studies 1970-1975 reported by
F. Fukuda (1977, p. 76). Collected 27 Apr.
1982 by N. Agetsuma; PRIKU, 1 (skeleton
only). Collected 24 May 1982 by PRIKU staff;
PRIKU, 1 (skeleton only). NE80.
Kisuki-cho, Ohara-gun ( Ai^lP7K;^fflI); Shi-
mane, HONSHU; ca. 35°17'N, 132°54'E;
mtDNA samples collected before 2003 by Y.
Kawamoto (2002, p. 60). SW59.
Kitakomachi, Kamogawa-city (fIJII rf]db''J''fflI);
Chiba, HONSHU; 35°07'N, 140°03'E; col-
lected 7 and 8 Feb. 1998 by PRIKU staff;
PRIKU, 2 (skeletons only). NE125.
Kitamatasanrin, Nosegawa-mura, Yoshino-gun
(^i?IPi?MJIIWdbflS:ajW); Nara, HONSHU;
34°07'N, 135°39'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 35: XXI- 1). See Fig. 2 A.
Kitamine, Kinsha-mura, Uma-gun {^MW>^^
^ it ^ ); Ehime, SHIKOKU; 33°54'N,
133°34'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 18:IX-11). SeeFig. 2A.
Kitamura, lijima-machi, Kamiina-gun (_b#BP?P
tSSfflJdb^); Nagano, HONSHU; 35°40'N,
137°54'E; collected 31 Oct. and 2 Dec. 1997
by M. Aimi; PRIKU, 1 1 (skeletons only, in-
cluding 4 with fragmented skulls and 1 with
postcranials only). NE77.
Kitanada-son, Itano-gun (^if^Pdbil^^); Toku-
shima, SHIKOKU; 34°12'N, 134°31'E; re-
ported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 24:
XI-34). See Fig. 2A.
Kitanokawa vicinity, Koguchi-mura, Higashi-
muro-gun (^^ftlPd^ P^dby JHftfe); Wa-
kayama, HONSHU; 33°45'N, 135°52'E; re-
ported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 34:
XX-48). See Fig. 2A.
Kitayama, Shioze-mura, Arima-gun (W/^?P^
)i ^ Jb UJ ); Hyogo, HONSHU; 34°48'N,
135°18'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 30: XVIII-29). See Fig. 2A.
Kitousanrin, Kamikitou-son, Kaifu-gun (^qP?P
±7KM^^7K1IUj^); Tokushima, SHIKOKU;
33°46'N, 134°14'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 23: Xl-4). See Fig. 2A.
Kitousanrin, Okukitou-son, Kaifij-gun (^nP?P
||7KlI;^7KIIli|#); Tokushima, SHIKOKU;
33°45'N, 134°10'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 23: XI-3). See Fig. 2A.
Kiwadahata, Kimitsu-city (^/$ Tti H^ BB !lffl);
Chiba, HONSHU; 35°11'N, 140°0rE; col-
lected Jul. 1996-Dec. 1998 by PRIKU staff;
PRIKU, 6 (skeletons only, including 1 with
fragmented skull). NE120.
Kiyano vicinity, Ookawa-mura, Tosa-gun (zhfe
IPAJIl^TKMifflfe); Kochi, SHIKOKU;
33°48'N, 133°30'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 22: X-29). See Fig. 2A.
Kiyokawa-mura, Aikou-gun ( S ¥ ?P /ff J 1 1 W );
Kanagawa, HONSHU; ca. 35°28'N, 139°17'E;
reported present before 1978 by Study Group
FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES
145
on the Present Status of Japanese Monkeys
(1977b, p. 23). MtDNA samples collected be-
fore 1998 by Y. Kawamoto (1997, p. 33; 2002,
p. 60). NEWS.
Kiyosumiyama (/^Mlll) (= Motokiyosumiyama;
Mt. Kiyosumi); Chiba, HONSHU; ca. 35°08'N,
140°12'E; reported before 1965 by K. Ka-
wanaka (1973, p. 115). Troop distribution
mapped by A. Suzuki (1972, p. 334). NE125.
Ko. See Kojiba.
Kobe-city (ttpTtT); Hyogo, HONSHU; 34°41'N,
135°10'E, reported origin of monkey obtained
2 Aug. 1904 by National Zoological Park,
Washington, D.C.; USNM (specimen received
12 Jun. 1912), 1. Not mapped.
Kochi-cho, Kamo-gun ( M j^ ?P M 1*1 fflj ); Hi-
roshima, HONSHU; 34°28'N, 132°53'E; pro-
visoned group; birth season, 1958-1959, re-
ported by Kawai et al. (1967, pp. 37, 39).
Blood samples collected before 1992 by No-
zawa et al. (1991, p. 414; 1996, p. 6). SW63.
Koganezawa vicinity, Nanaho-mura, Kita-
tsuru-gun (db|[PlgiP-b^^d^^)Rftfe); Ya-
manashi, HONSHU; 35°41'N, 138°57'E; re-
ported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 48:
XXXIII-25). See Fig. 2A.
Kogarashigataniyama vicinity, Sawatani-mura,
Oochi-gun (E^?P)i#W7Ktt#ajffe); Shi-
mane, HONSHU; 35°05'N, 132°38'E; reported
in questionnaire survey conducted in 1923 by
K. Hasebe (Iwano, 1974, p. 28: XVI-26). See
Fig. 2A.
Kohoku-cho, Higashiazai-gun (^/^#?P/Sdb
BU); Shiga, HONSHU; ca. 35°26'N, 136°15'E;
mtDNA samples collected before 2003 by Y.
Kawamoto (2002, p. 60). SW6.
Koide, Kuramata-mura, Nakauonuma-gun (4^^
>S iP 5t ^ W ^J^ lii ); Niigata, HONSHU;
36°58'N, 138°44'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 47: XXXlI-2). See Fig. 2A.
Kojiba, Kamiyaku-cho, Kumage-gun (^^?P_h
WX^HM^y, Kagoshima, YAKUSHIMA;
ca. 30°22'N, 130°23'E; observed 1975-1979
by T. Maruhashi (1982, p. 318; cf. Takasaki &
Masui, 1984, p. 311). SfF/^y.
Kojima, Ichiki-son, Minaminaka-gun (j^SPlRliP
TfT^KW^fi); Miyazaki, KOJIMA; 31°27'N,
131°23'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 15:11-3). See Fig. 2A.
Kojima, Kushima-city ($f^r(i^A) (= Ko-
shima); Miyazaki, KOJIMA; 3r27'N,
131°23'E; reported present before 1939 by N.
Kuroda (1938, p. 112; 1940, p. 271). Provi-
sioning history, 1952-1977, reported by A.
Mori (1979b, p. 372). Birth season, 1956-1966,
reported by Kawai et al. (1967, p. 39). Col-
lected 11 Sep. 1969-29 Sep. 1996 by KFL
staff; KFL, 48 ( 1 skin only; 47 skeletons only,
including 3 with fragmented skulls, 8 with
missing mandibles, and 1 with missing cra-
nium). Blood samples collected before 1992 by
Nozawa et al. (1991, p. 414; 1996, p. 6). Mi-
crosatellite DNA analyzed before 1996 by
Domingo-Roura et al. (1997, p. 358). External
measurements taken before 1997 by Hamada
et al. (1996a, pp. 98, 99). MtDNA samples
collected before 2003 by Y. Kawamoto (2002,
p. 60). SW97.
Koka-cho, Koka-gun (¥M^¥MfflI); Shiga,
HONSHU; ca. 34°54'N, 136°14'E; mtDNA
samples collected before 2003 by Y. Kawa-
moto (2002, p. 60). 5^/7.
Kokurano, Ootaki-machi, Isumi-gun (MPfI^A
^^fflJ/JN^tif); Chiba, HONSHU; 35°12'N,
140°09'E; collected 10 Aug. and 8 Sep. 1997
by PRIKU staff; PRIKU, 3 (skeletons only).
NE126.
Komagane Mountain Range, Hiyoshi-mura, Ni-
shichikuma-gun (M^^iPSM^ffil'^-^fgUj
M); Nagano, HONSHU; 35°51'N, 137°47'E;
reported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 46:
XXXI-10). SeeFig. 2A.
Komagatake, Kanegasaki-mura, Isawa-gun (ll/R
^^T^^^T^); hvate, HONSHU;
39°irN, 141°06'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 56: XLV-7). See Fig. 2A.
Komagatake Mountain Range, Agematsu-machi,
Nishichikuma-gun (ffimSIP±^A^fflJ|fi|<r§
UjM); Nagano, HONSHU; 35°46'N, 137°42'E;
reported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 45:
XXXl-[6]). See Fig. 2A.
Komagatake Mountain Range, Sugawara-mura,
146
FIELDIANA: ZOOLOGY
Kitakoma-gun (db&J5?PWi^W|fi)^l&UjM);
Yamanashi, HONSHU; 35°48'N, 138°18'E;
reported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 48:
XXXIII-17). SeeFig. 2A.
Komagatake vicinity, Nechi-mura, Nishi-
kubiki-gun (mm^W^^^^^T ^^)- Nii-
gata, HONSHU; 36°57'N, 137°53'E; reported
in questionnaire survey conducted in 1923 by
K. Hasebe (Iwano, 1974, p. 47: XXXII- 1). See
Fig. 2A.
Komagatake vicinity, Takane-mura, Iwafune-gun
{^^^Wmm-^^T ^^)- Niigata, HONSHU;
38°20'N, 139°37'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 48: XXXII-23). See Fig. 2A.
Komataoku vicinity, Asuka-mura, Minami-
muro-gun (j^^ftl^miWd^J^^ftfe); Mie,
HONSHU; 33°57'N, 136°07'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 37: XXIII-50). See
Fig. 2A.
Komatsu-mura, Shiga-gun ( ^^ M ?P d^ ^A^ ^^ );
Shiga, HONSHU; 35°15'N, 135°57'E; reported
in questionnaire survey conducted in 1923 by
K. Hasebe (Iwano, 1974, p. 38: XXIV-5). See
Fig. 2A.
Komatsushima-cho and Ikuhina-son, Katsu-
ura-gun {Bm^^l^^m^ il^ttP^m To-
kushima, SHIKOKU; 33°55'N, 134°28'E; re-
ported absent in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 23: XI- 17). Not mapped.
Komeyama, Noda-mura, Kariwa-gun (X'J^^lPi?
H ;f^Nt tI^ Uj ); Niigata, HONSHU; 37°16'N,
138°34'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 47: XXXII- 12). SeeFig. 2A.
Komogawa, Asahi-mura, Iwafune-gun (^^p^
^S+^SJII); Niigata, HONSHU; 38°18'N,
139°36'E; collected 5 Oct. 1998 by M. Aimi;
PRIKU, 1 (skeleton only). Collected 23 Oct.
1998 by PRIKU staff; PRIKU, 2 (skeletons
only). NE31.
Komori, Nishiyama-mura, Minamimuro-gun (^
#«lPSUj4^d^^); Mie, HONSHU; 33°54'N,
135°58'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 37: XXIII-53). See Fig. 2A.
Konahisanrin vicinity, Nishiwara-mura,
Gujo-gun (l^±^SS]MWd^J§Pi;baj^ftfe);
Gifu, HONSHU; 35°41'N, 137°02'E; reported
in questionnaire survey conducted in 1923 by
K. Hasebe (Iwano, 1974, p. 41: XXVIII-24).
See Fig. 2A.
Konan National Forest vicinity, Ashio-machi,
Kamitsuga-gun (±i5S?PS^fflT/^]tSW^
fte); Tochigi, HONSHU; 36°36'N, 139°27'E;
reported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 51:
XXXIX-2). See Fig. 2A.
Kongoumine vicinity, Mikawa-mura, Nishi-
muro-gun (M#«?PHjl|;^#ilJ^fte); Wa-
kayama, HONSHU; 33°40'N, 135°35'E; re-
ported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 34:
XX-27). See Fig. 2A.
Konishi, Kaita-mura, Kiso-gun (^eiP^BB^
^J^M); Nagano, HONSHU; 35°59'N, 137°35'E;
collected 23 Aug. 1998 by M. Aimi; PRIKU, I
(skeleton only). NE85.
Konishioku, Nishino-son, Mitsugi-gun (^H?P
M i? ^ ^J^ S H ); Hiroshima, HONSHU;
34°25'N, 133°03'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 26: XIV-26). See Fig. 2A.
Konoshiro, Takano-mura, Takeno-gun ("friJ^P
t^ i? ^^ jib -ft ); Kyoto, HONSHU; 35°43'N,
135°08'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 35:XXII-1). SeeFig. 2A.
Koryujiyama, Kamisano-mura, Kumano-gun {M.
i?15±^i)l;|4iSfl#aj); Kyoto, HONSHU;
35°33'N, 135°00'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 35: XXII-7). See Fig. 2A.
Kosadake, Higashitomochi-mura, Shimo-
mashiki-gun (TM^lPm^iKffl W¥feS);
Kumamoto, KYUSHU; 32°38'N, 130°56'E;
reported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 16: IV-8).
See Fig. 2A.
Koshikiya, Ichihara-city {'^W^'^'^%/^); Chiba,
HONSHU; 35°20'N, 140°10'E; collected 5 Feb.
1999 by PRIKU staff; PRIKU, 1 (skeleton
only). NE127.
Koshima. See Kojima.
Koshima, Yaku-cho, Kumage-gun (S^^^M^
FOODEN AND AIMI: SYSTEMATIC REVIEW OE JAPANESE MACAQUES
147
fflld^l); Kagoshima, YAKUSHIMA; 30°14'N,
130°32'E; collected 28 Jul. 1987 by PRIKU
staff; PRIKU, 1 (skeleton only, with frag-
mented skull). SW102.
Kouchi, Yokokura-mura, Ibi-gun (SHIPHIMW
Z\0^); Gifu, HONSHU; 35°32'N, 136°36'E;
reported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 40:
XXVIIl-9). See Fig. 2A.
Koujiro, Mitoya-cho, lishi-gun (^^?PHZJM
BT # -ft ); Shimane, HONSHU; 35°13'N,
132°53'E; collected 30 Nov. and 9 Dec. 1994
by PRIKU staff; PRIKU, 3 (skeletons only, in-
cluding 1 with fragmented skull and 2 with
postcranials only). SW59.
Kouwayama vicinity, Suhara-mura, Mugi-gun
{^%W,^W^WM^\^^)\ Gifu, HONSHU;
35°36'N, 136°58'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 41: XXVIII-18). See Fig. 2A.
Koyadaira, Koyadaira-son, Oe-gun (^^IP^M
¥ ^ ^ M ¥ ); Tokushima, SHIKOKU;
33°54'N, 134°irE; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 23: XI-28). See Fig. 2 A.
Koyayama, Ukena-mura, Kamiukena-gun (_L/?
Tv: ?P )? A ^ ^h Jl OJ ); Ehime, SHIKOKU;
33°30'N, 132°52'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 21: IX-22). See Fig. 2 A.
Kozagawa-cho, Higashimuro-gun (^#- ft IP "4"
MJIIfflJ); Wakayama, HONSHU; ca. 33°32'N,
135°48'E; mtDNA samples collected before
2003 by Y. Kawamoto (2002, p. 60). SW28.
Kozawamata, Ootaki-machi, Isumi-gun (MP^?P
A^*fflT^h)RX); Chiba, HONSHU; 35°13'N,
140°10'E; collected 23 Oct. 1998-25 Jan. 1999
by PRIKU staff; PRIKU, 4 (skeletons only, in-
cluding 2 with fragmented skulls). NE126.
Kuchisubosanrin, Hase-mura, linan-gun (^j^lP
1^M¥iU^\h^)\ Mie, HONSHU; 34°25'N,
136°10'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 37: XXIII-22). See Fig. 2A.
Kuchiyama-son, Mima-gun ( II /^ IP P ill W );
Tokushima, SHIKOKU; 34°01'N, 134°09'E;
reported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 23:
XI-24). See Fig. 2A.
Kugayama National Forest vicinity, Yamana-son,
Fukayasu-gun (J^^^Oji^W^MUj 1^^
j\k); Hiroshima, HONSHU; 34°40'N, 133°23'E;
reported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 26:
XIV-31). SeeFig. 2A.
Kugino-mura, Aso-gun {MM^X^^H); Kii-
mamoto, KYUSHU; ca. 32°48'N, 13r01'E;
mtDNA samples collected before 2003 by Y.
Kawamoto (2002, p. 60). SW88.
Kuigautsu, Hinui-mura, Oochi-gun (B^lPBm
^^TO'V); Shimane, HONSHU; 34°53'N,
132°23'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 28:XVI-15). SeeFig. 2A.
Kujuu vicinity, Nakasuji-mura, Hata-gun (iRI^
^^Wi^%^^); Kochi, SHIKOKU;
32°57'N, 132°50'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 21: X-9). See Fig. 2 A.
Kujuu-mura/Tamakikuchi-mura, Higashi-
muro-gun (^^S1P:XM^5^P^); Wa-
kayama, HONSHU; 33°51'N, 135°5rE; re-
ported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 34:
XX-53). SeeFig. 2A.
Kumabushiyama, Yaekawachi-mura, Shi-
moina-gun {TWMW>J\W.nP^^m^\^);
Nagano, HONSHU; 35°15'N, 137°53'E; re-
ported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 47:
XXXI-51). SeeFig. 2A.
Kumadayama, Nakamata-mura, Iwafline-gun (^
^p' ^ 4^ ^M ^ ^ ea UJ ); Niigata, HONSHU;
38°3rN, 139°36'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 48: XXXII-26). See Fig. 2A.
Kumamidani vicinity, Kuroda-mura, Kitaku-
wada-gun (db^ffllPHBa^l^m^ffe); Kyoto,
HONSHU; 35°13'N, 135°44'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 36: XXII- 19). See Fig.
2A.
Kumano-city {MWt'^); Mie, HONSHU; ca.
33°53'N, 136°07'E; mtDNA samples collected
before 2003 by Y. Kawamoto (2002, p. 60).
SW25.
Kumayama-cho, Akaiwa-gun (55^^?Pfl^li| fff);
Okayama, HONSHU; 34°47'N, 134°05E;
148
FIELDIANA: ZOOLOGY
mtDNA samples collected 1998-2000 by I.
Yoshimi and H. Takasaki (2003, p. 71). SW52.
Kunimi vicinity, Higashinakasuji-mura,
Hata-gun i^^mM^M^mM.i\k); Kochi,
SHIKOKU; 32°57'N, 132°52'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 21: X-5). See Fig.
2A.
Kunimidaira vicinity, Uchinoura-mura, Ki-
motsuki-gun (flf M ^ 1*1 ;^M^ @ .^^ftfe);
Kagoshima, KYUSHU; 31°15'N, 131°03'E;
reported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 12: 1-16).
See Fig. 2A.
Kuniwaridake (= Mt. Kuniwari), Kamiyaku-cho,
Kumage-gun (^.^^±M^fflT@SlJS); Ka-
goshima, YAKUSHIMA; 30°20'N, 130°25'E;
collected 27 Jul. 1981 by PRIKU staff; PRIKU,
1 (skeleton only, with fragmented skull).
SWIOL
Kuniwaridake (= Mt. Kuniwari), southeastern
slopes, Kamiyaku-cho, Kumage-gun (SI^^P
±M^ffliaflJ^I^K#4®), ca. 900-1200 m
elevation; Kagoshima, YAKUSHIMA; ca.
30°20'N, 130°26'E; ca. 11 troops censused
Jul.-Aug. 1995 by S. Yoshihiro et al. (1999, p.
4\\).SWW1.
Kuniwaridake (= Mt. Kuniwari), summit area,
Kamiyaku-cho, Kumage-gun (^l^^-hM^
fflJSitJS]I±), ca. 1200-1323 m elevation;
Kagoshima, YAKUSHIMA; ca. 30°21'N,
130°25'E; 4 troops censused Jul.-Aug. 1995
by S. Yoshihiro et al. (1999, p. 41 1). SWIOI.
Kuniwaridake (= Mt. Kuniwari), western slopes,
Kamiyaku-cho, Kumage-gun (^^^-hM^
ffllSilJSffif^®), ca. 300-900 m elevation;
Kagoshima, YAKUSHIMA; ca. 30°20'N,
130°24'E; ca. 13 troops censused Jul.-Aug.
1995 by S. Yoshihiro et al. (1999, p. 411).
DNA samples collected before 2005 by X.
Domingo-Roura et al. (2004, p. 33). Not
mapped.
Kuniyama National Forest, Iwanashi-son,
Akaiwa-gun ('^^^S^WSUjSW^);
Okayama, HONSHU; 34°50'N, 134°09'E; re-
ported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 27:
XV-20). See Fig. 2A.
Kuradama, Kimitsu-city (^/^rtTMi); Chiba,
HONSHU; 35°13'N, 140°07'E; collected 2 Jul.
1996-5 Mar. 1999 by PRIKU staff; PRIKU,
31 (skeletons only, including 3 with frag-
mented skulls). NE126.
Kuragariyama, Kamiyama-mura, Uma-gun (^S
W±lh^'y'7:^U[Uy, Ehime, SHIKOKU;
33°58'N, 133°4rE; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 21: IX- 13). See Fig. 2A.
Kuraishidake, Miya-mura, Katsuta-gun (XlJBQ^P
^^^^11); Miyagi, HONSHU; 38°0rN,
140°37'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 55:XLIII-4). SeeFig. 2A.
Kurase vicinity, Sakuragi-mura, Shusou-gun (M
mW^^^^Mi\^); Ehime, SHIKOKU;
33°49'N, 133°0rE; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 18: IX-7). See Fig. 2A.
Kuratani vicinity, Kitagouchi-son, Kuga-gun (^
iRl ?P db )rI rt W ^ ^ ffe ); Yamaguchi,
HONSHU; 34°09'N, 132°05'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 25: XIII- 13). See Fig.
2A.
Kuratani vicinity, Saigawa-mura, Ishikawa-gun
(^JIIiPJlJIW5t#ffe); Ishikawa, HONSHU;
36°28'N, 136°42'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 39: XXVI- 1). See Fig. 2A.
Kurataniyama, Tataragi, Nakagawa-mura,
Asago-gun im^W>^)\m^^ ^^^t'^lh);
Hyogo, HONSHU; 35°15'N, 134°49'E; re-
ported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 30:
XVIII-15). SeeFig. 2A.
Kurio-Segire. See Yakushima, Census Area 6.
Kurisuyama, Joto-mura, Niwa-gun (^^?Pi^K
^ ^ ffi Uj ); Aichi, HONSHU; 35°21'N,
137°00'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 40: XXVII-9). See Fig. 2A.
Kuriyama-mura, Shioya-gun (^#^^li|^);
Tochigi, HONSHU; ca. 36°54'N, 139°35'E;
collected 2 Aug. 1972 and 18 Sep. 1986 by
TPM staff; TPM, 2 (skulls only, including 1
with mandible missing). NE42.
Kurobe (H^); Toyama, HONSHU; 36°39'N,
137°41'E; observed Nov. 1979-Jan. 1987 by H.
FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES
149
Akaza(1988,p. \2).NE54.
Kurobedani,Uchiyama-mura, Shimoniikawa-aun
( T Sf Jll ?P rt OJ ^ H ^ ^ ); Toyama,
HONSHU; 36°51'N, 137°32'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 42: XXIX-9). See Fig.
2A.
Kuroda vicinity, Takane-son, Kuga-gun (3Eft.iRllP
R S ^ H ea -te ); YamaguchC HONSHU;
34°24'N, 132°03'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 25: XIII- 14). See Fig. 2 A.
Kurohimeyama, Takayanagi-mura, Kariwa-gun
(XlJ^?PS^^IIieaj);^M/ga/fl, HONSHU;
37°11'N, 138°38'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 47: XXXII- 13). See Fig. 2A.
Kurohimeyama, Ukawa-mura, Kariwa-gun (X'J^
^ 56IJII ^ II ie UJ ); Niigata, HONSHU:
37°13'N, I38°34'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 47: XXXII- 1 1). See Fig. 2 A.
Kurokagara vicinity, Asuna-mura, Oochi-gun (e
^W>n^M^W.ti1i^ ^)\ Shimam.
HONSHU; 34°53'N, 132°37'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 28: XVI-20). See Fig.
2A.
Kurokawa, Hirata-mura, Hata-gun (IS^IP^BQ
^ H Jll ); Kochi, SHIKOKU; 32°56'N.
132°49'E; reported in questionnaire sur\ey
conducted in 1 923 by K. Hasebe (Iwano, 1 974.
p. 21:X-4). SeeFig. 2A.
Kurokawa-mura, Kitakanbara-gun (dbSJ^lPM
Jll ^ ); Niigata, HONSHU; ca. 38°04'N.
I39°26'E; mtDNA samples collected before
1998 by Y. Kawamoto (1997, p. 33; 2002. p.
60). NE33.
Kurokawayama, Awano-mura, Tsuruga-gun (^
MIPHS^H^rTOJ); FukuL HONSHU;
35°34'N, 136°03'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 39: XXV- 16). See Fig. 2 A.
Kuromoriyama vicinity. Nishiokuni-mura,
Mogami-gun ( S ± IP ® ^M ^hTH ^ Oj ffe );
Yamagata, HONSHU; 38°48'N, 140°29'E; re-
ported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 55:
XLII-10). See Fig. 2 A.
Kurosawa National Forest. Ogura-mura, Mi-
namiazumi-gun (]^$^iP'J^li"^M/RB W
^); Nagano, HONSHU; 36°20'N, 137°50'E;
reported in questionnaire sur\'ey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 46:
XXXI-16). SeeFig. 2A.
Kurosu vicinity. Shimazu-mura, Watarai-gun (]^
^^Pft^^H^-te); Mie, HONSHU; 34°15'N,
136°27'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 37:XXIII-31). SeeFig. 2A.
Kurotaki. Yamauchi-mura, Koka-gun (^MlPill
1*1 ^ II )i ); Shiga, HONSHU; 34°55'N,
136°20'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 38: XXIV-26). See Fig. 2A.
Kurouchi, Kiso-mura, Kiso-gun (^e^^ffi^
Urt); Nagano, HONSHu'J 35°58'N. 137°45'E;
collected 12 Dec. 1997 by M. Aimi; PRIKU, I
(skeleton only). NE86.
Kusaki-mura, Sayo-gun (■feffl^P^i'^W); Hyogo,
HONSHU; 34°56'N. 134°18'E; reported in
questionnaire sur\'ey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 30: XVIII-3). See Fig.
2A.
Kusenbuyama vicinity. Minamihata-mura. Chi-
kushi-gun {m^^^m^h=f^\h^): Fu-
kuoka, KYUSHU; 33°24'N, 130°26'E; reported
in questionnaire survey conducted in 1923 by
K. Hasebe (Iwano, 1974, p. 17: VII- 1). See Fig.
2A.
Kushigawa. Kaifu-cho, Kaifu-gun (^pPlP^qP
urr^iJII); Tokushima, SHIKOKU; 33°36'N,
134°17'E; collected 19 Sep. 2001 by M. Aimi;
PRIKU, \.SW73.
Kushigawa vicinity. Kawanishi-son, Kaifu-gun
( m^ ^ Jll S ^ # Jll ^ ); Tokushima,
SHIKOKU; 33°35'N. 134°17'E; reported in
questionnaire sur\ey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 23: XI- 1). See Fig.
2A.
Kushima-city (^ralrfJ); Miyazaki. KYUSHU; ca.
3^287^, 131°12'E; blood samples collected
before 1992 by Nozawa et al. (1991, p. 414;
1996, p. 6). MtDNA samples collected before
2003 by Y. Kawamoto (2002, p. 60). SW96.
Kusoudomari, Wakinosawa-mura. Shi-
mokita-gun {Ti^^^WR^XB.^); Ao-
mori, HONSHU; 41°09'N, 140°47'E; found
150
FIELDIANA: ZOOLOGY
and donated in 1964 by unknown collector (cf.
Study Group on the Present Status of Japanese
Monkeys, 1977a, p. 10, locality Aomori-2);
JMC, 1 (skull only). NE5.
Kusugawa, Kamiyaku-cho, Kumage-gun (^^
IP ± Jl ^ ffll ffi Jll ); Kagoshima,
YAKUSHIMA; 30°23'N, 130°35'E; collected
25 Jul. 1987 by PRIKU staff; PRIKU, 1
(skeleton only). SWIOO.
Kuwashimachinai, Shiramine-mura, Nomi-gun
( tg H ?P a III* ^ # ft itfe I*] ); Ishikawa,
HONSHU; 36°10'N, 136°39'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 40: XXVI-6). See Fig.
2A.
Kuwasononanbu, Matsukawa-machi, Shi-
moina-gun (T^IPlP^AMIIffll^Ult^); Na-
gano, HONSHU; 35°36'N, 137°52'E; collected
11 Dec. 1997 by M. Aimi; PRIKU, 4 (skele-
tons only). NE76.
Kyogatake ( i^ "^ S ); Nagasaki-Saga border,
KYUSHU; ca. 32°59'N, 130°04'E; observed in
1940s by local residents (Sakura, 1976, p. 151);
subsequently extinct at this locality (Ikeda and
Eguchi, 1978, p. 59). 5^(97.
Kyonan-machi, Awa-gun (^J^^PISl^fflJ);
Chiba, HONSHU; ca. 35°06'N, 139°52'E;
mtDNA samples collected before 1999 by Y.
Kawamoto (1998, p. 54; 2002, p. 60). NEI20.
Kyouhou vicinity, Hajikami-mura, Hata-gun (♦#
^?P^±^^t:^)iffe); Kochi, SHIKOKU;
33°00'N, 132°43'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 21: X-1 1). See Fig. 2 A.
Launceston, Tasmania; 4r35'S, 147°22'E;
translocation in 1980 from Kasuga, Hyogo
Prefecture, reported by M. Nozaki (1993, p.
j97). Not mapped.
Maemoriyama vicinity, Higashiokuni-mura,
Mogami-gun (ft ±1^^^^ S ^lu I^Ul ffe);
Yamagata, HONSHU; 38°46'N, 140°33'E; re-
ported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 55:
XLIl-11). SeeFig. 2A. ^
Maki, Takayama-mura, Kamitakai-gun (Jiisi^
^PilUj;^!"^); Nagano, HONSHU; 36°40'N,
138°25'E; collected 19 Jul. 1998 by M. Aimi;
PRIKU, 1 (skeleton only). NE49.
Makino-cho, Takashima-gun (i^ S?P'7 ^ y BJ),
Shiga, HONSHU; 35°30'N, 136°03'E; ob-
served Aug.-Sept. 1989 by M. Nakamichi et al.
(Yamagiwa & Hill, 1998, p. 267). SW3.
Makinosaki vicinity, Oohara-mura, Oshika-gun
(ttElPAJ^^^(7)llif ffe); Miyagi, HONSHU;
38°2rN, 141°30'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 55: XLIII-1 1). See Fig. 2A.
Makiosan vicinity, Makinoshima-mura,
Kuze-gun (r^^iP^ft^^mUjftfe); Kyoto,
HONSHU; 34°53'N, 135°46'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 36: XXII-28). See Fig.
2A.
Makitaniyama vicinity, Umaji-mura, Aki-gun (^
^^m^^W^\h^); Kochi, SHIKOKU;
33°32'N, 134°05'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 22: X-39). See Fig. 2A.
Mantaroyama vicinity, Tsuchidaru-mura, Mi-
namiuonuma-gun (]^^5S?P±#^ll:^l!Plil
fte); Niigata, HONSHU; 36°53'N, 138°52'E;
reported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 47:
XXXII-4). See Fig. 2A.
Manzayama, Tsumagoi-mura, Agatsuma-gun (^
^ IP If i2^x ^ 77 0 111 ); Gunma, HONSHU;
36°38'N, 138°30'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 51: XL-8). See Fig. 2 A.
Maruseyama, Ichigi-mura, Oochi-gun (e^^Tfi"
:^¥iSlM\^); Shimane, HONSHU; 34°51'N,
132°25'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 28: XVI- 16). SeeFig. 2A.
Maruyama, Ubuchi-mura, Usui-gun (fltt^XlP/^
^J ^ ^ UJ ); Gunma, HONSHU; 36°25'N,
138°46'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 51:XL-6). SeeFig. 2A.
Masakiyama, Ipponmatsu-mura, Minamiuwa-gun
(jt^^^P-^^i^IETKUj); Ehime,
SHIKOKU; 32°59'N, 132°40'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 18: IX-4). See Fig.
2A.
Masakiyama, Tsutsuga-son, Yamagata-gun (|JL|
!f;15MM^IE7KUj); Hiroshima, HONSHU;
34°35'N, 132°15'E; reported in questionnaire
FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES
151
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 26: XIV-4). See Fig. 2A.
Masezawa, Hachimori-mura, Yamamoto-gun (llj
^W>J\M^MM'yR); Akita, HONSHU;
40°22'N, 140°07'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 56: XLIV-9). See Fig. 2A.
Masudayama, Nakatane-mura, Kumage-gun {M.
^W^U^^i^miU); Kagoshima,
TANEGASHIMA; 30°35'N, 130°59'E; re-
ported as extinct in questionnaire survey con-
ducted in 1923 by K. Hasebe (Iwano, 1974, p.
12:1-3). See Fig. 2A.
Masugata, Takayama-mura, Kamitakai-gun (Ji
mk^mih^mM); Nagano, HONSHU;
36°4rN, 138°22'E; collected 7 Oct. and 1 Nov.
1998 by M. Aimi; PRIKU, 3 (skeletons only).
NE49.
Matani, Agira-mura, Kawage-gun (5nJS^Q^:|4
M-^y, Mie, HONSHU; coordinates unknown;
reported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 38:
XXIII-58). Not mapped.
Matsubakawayama vicinity, Matsubakawa-mura,
Takaoka-gun (I^I?^?P^A^SJI|^^iSJI| UUffe);
Kochi, SHIKOKU; 33°16'N, 133°04'E; re-
ported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 22: X-19).
See Fig. 2A.
Matsukasa-mura, lishi-gun ( tS S ?P ^ S ^ );
Shimane, HONSHU; 35°11'N, 132°42'E; re-
ported (as possibly temporary habitat) in ques-
tionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 28: XVI-33). See Fig.
2A.
Matsuno-cho, Kitauwa-gun (db^^?P^ii?fflI);
Ehime, SHIKOKU; ca. 33°12'N, 132°44'E;
mtDNA samples collected before 2003 by Y.
Kawamoto (2002, p. 60). SW78.
Mayasan, Nishinada-mura, Muko-gun (K0^S
li;^^t0gP^J); Hyogo, HONSHU; 34°44'N,
135°12'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 30: XVIII-30). See Fig. 2A.
Mera, Tateyama-city ( tl ill TtT ^ ^ ); Chiba,
HONSHU; 34°54'N, 139°49'E; collected 12
Jul. 1997 by PRIKU staff; PRIKU, 1 (skeleton
only). NE122.
Metaka, Ooshika-mura, Shimoina-gun (T#3P
^XU¥it:m)\ Nagano, HONSHU; 35°38'N,
138°03'E; collected 31 Dec. 1997 by M. Aimi;
PRIKU, 1 (skeleton only). NE70.
Michikata vicinity, Nakajima-mura, Watarai-gun
(Jt^^4^ft^]l:^'te); Mie, HONSHU;
34°18'N, 136°35'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 37: XXIII-37). See Fig. 2A.
Mie ( H M ) (= Miye), HONSHU;
33°44'-35°15'N, 135°52'-136°54'E; collected.
4 Jan. 1939 by T. Barbour; MCZ, 1. Not
mapped.
Mihama-cho, Mikata-gun {=.^5^^)^^); Fu-
kiti, HONSHU; ca. 35°36'N, 135°56'E;
mtDNA samples collected before 2003 by Y.
Kawamoto (2002, p. 60). SW2.
Mihama-cho, Minamimuro-gun (l^-^ft^P^^
fflj); Mie, HONSHU; ca. 33°48'N, 136°02'E;
mtDNA samples collected before 2003 by Y.
Kawamoto (2002, p. 60). SW26.
Mihama-mura, Kamo-gun (Mj^^PH^^); Shi-
zuoka, HONSHU; 34°43'N, 138°46'E; reported
in questionnaire survey conducted in 1923 by
K. Hasebe (Iwano, 1974, p. 45: XXX-24). See
Fig. 2A.
Mihara-city (Hj^TtT); Hiroshima, HONSHU;
34°24'N, 133°05'E; external measurements
taken before 1997 by Hamada et al. (1996a, pp.
98, 99). SW64.
Mikagura, Nishikawa-mura, Higashikanbara-gun
(^aj^?PMJI|^1itt2$<); Niigata, HONSHU;
37°37'N, 139°28'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 47: XXXII-16). See Fig. 2A.
Mikata-cho, Mikata-gun (H;?31PH;J3 fflJ); Fukui,
HONSHU; 35°37'N, 135°51'E; captured 1
May 1968 by JMC staff, died in captivity 24
May-5 June 1968; JMC, 3 (skulls only). Blood
samples collected before 1992 by Nozawa et al.
(1991, p. 414; 1996, p. 6).5Pr7.
Mikatasanrin, Ya-mura, Mikata-gun (H^?PA
;^^t H :i^ Oj :|9^ ); Fukui, HONSHU; 35°33'N,
135°55'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 39:XXV-13). SeeFig. 2A.
Mikoshi vicinity, Misato-mura, Higashimuro-gun
(K^«?PHM^H^^); Wakayama,
HONSHU; 33°51'N, 135°43'E; reported in
questionnaire survey conducted in 1923 by K.
152
FIELDIANA: ZOOLOGY
Hasebe (Iwano, 1974, p. 34: XX-52). See Fig.
2A.
Mikuniyama, Takahashi-mura, Izushi-gun (t±i5
iPilM^HSUj); Hyogo, HONSHU;
35°26'N, 135°0rE; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 30: XVIII-20). See Fig. 2 A.
Mikusu, Minakami-cho, Ooda-city (AB9 rfT7K_L
ffllH:^^); Shimane, HONSHU; 35°04'N,
132°26'E; collected 15 Aug. 1994 by PRIKU
staff; PRIKU, 4 (skeletons only). SW6L
Mimaisan vicinity, Mimai-mura, Nishimuro-gun
(ffi^ftfPH^^^H^Ojffe); Wakayama,
HONSHU; 33°36'N, 135°29'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 34: XX-29). See Fig.
2A.
Minamikawa vicinity, Mori-mura, Tosa-gun (i
te ^ 1^ ^ it Jll ^ ); Kochi, SHIKOKU;
33°45'N, 133°33'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 22: X-30). See Fig. 2A.
Minamikomagadake. Nanakubo-mura, Kami-
ina-gun {^L^M^^zXU^'^^'r ^); Na-
gano, HONSHU; 35°39'N, 137°53'E; reported
in questionnaire survey conducted in 1923 by
K. Hasebe (Iwano, 1974, p. 46: XXXI-43). See
Fig. 2A.
Minamikurosawa National Forest, Azusa-mura,
Minamiazumi-gun (it$S?P#^^]tM/H@
W^); Nagano, HONSHU; 36°14'N, 137°50'E;
reported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 46:
XXXI-13). SeeFig. 2A.
Minamimachi, Suwa-city (Ml^rfTj^fflJ); Nagano,
HONSHU; 36°00'N, 138°08'E; collected 26
Jul. 1998 by M. Aimi; PRIKU, 1 (skeleton
only). NE95.
Minamitoyosawayama vicinity, Yukuchi-mura,
Hienuki-gun (Pml^^^P^lt^/RLljffe);
Iwate, HONSHU; 39°25'N, 141°06'E; reported
in questionnaire survey conducted in 1923 by
K. Hasebe (Iwano, 1974, p. 56: XLV-9). See
Fig. 2A.
Minoo-city ( ^ ® rf] ) (= Minou); Osaka,
HONSHU; ca. 34°51'N, 135°29'E; provisioned
group; birth season, 1957-1966, reported by
Kawai et al. (1967, pp. 37, 39). Collected 10
Apr. 1979 and 19 Jun. 1980 by PRIKU staff;
PRIKU, 3 (skeletons only). MtDNA samples
collected ca. 1986-1991 by Hayasaka et al.
(1991, p. 400). Blood samples collected before
1992 by Nozawa et al. (1991, p. 414; 1996, p.
6). MtDNA samples collected before 2003 by
Y. Kawamoto (2002, p. 60). SW53.
Minoosan National Forest, Minoo-mura,
Toyono-gun (MtelP^®^^® ill HW^);
Osaka, HONSHU; 34°49'N, 135°30'E; re-
ported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 33:
XIX-1). SeeFig. 2A.
Miokawa-mura, Higashimuro-gun (!^-^SlPH
MJII^); Wakayama, HONSHU; 33°34'N,
135°4rE; reported in questionnaire survey
conducted in 1 923 by K. Hasebe (Iwano, 1 974,
p. 34: XX-39). See Fig. 2A.
Misaki-Makinouchi vicinity, Toi-son, Minami-
naka-gun {"^MM^n^^imm^cn]^^)-
Miyazaki, KYUSHU; 31°23'N, 13r20'E; re-
ported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 12: II- 1).
See Fig. 2A.
Misogawa-goryorin vicinity, Kiso-mura, Ni-
shichikuma-gun (SmSlPTKia^D^PgJIIiai
^ W ftfe ); Nagano, HONSHU; 35°59'N,
137°46'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 46:XXXI-11). SeeFig. 2A.
Misokawa, Kiso-mura, Kiso-gun (^a^^ffi^t
^ Pi Jll ); Nagano, HONSHU; 35°58'N,
137°46'E; collected 30 Dec. 1997 by PRIKU
staff; PRIKU, 2 (skeletons only). NE86.
Mitakeyama, Aikawa-mura, Oono-gun (Ai??P
-^JHWISli^Lil); Oita, KYUSHU; 32°54'N,
131°30'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 16:111-5). SeeFig. 2A.
Mitakeyama-goryorin, Mitake-mura, Nishichi-
kuma-gun (ffimJSiPH|&;14|EPi^aj|i^#);
Nagano, HONSHU; 35°53'N, 137°36'E; re-
ported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 45:
XXXI-9). See Fig. 2A.
Mitoya-cho, lishi-gun (ISSlPHTJMffll); Shi-
mane, HONSHU; 35°17'N, 132°52'E; col-
lected 7 Feb. 1995-13 Mar. 1997 by PRIKU
staff; PRIKU, 5 (skeletons only, including 1
with fragmented skull; excluding 1 juvenile
FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES
153
skull mistakenly associated with data pertain-
ing to large adult male). SW59.
Mitsudake, Hata-mura, Taki-gun (^iS^^^H
it); Hyogo, HONSHU; 35°07'N, 135°14'E;
reported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 30:
XVIII-27). See Fig. 2A.
Mitsue-mura, Uda-gun (^PSIP^^^); Nara,
HONSHU; 34°30'N, 136°11'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 35: XXI-8). See Fig.
2A.
Mitsuminegawa upstream, Inasato-mura, Kami-
ina-gun (±^3PIP#SPM^H** Jl| ±)^);
Nagano, HONSHU; 35°41'N, 138°07'E; re-
ported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 46:
XXXI-44). See Fig. 2A.
Mitsutouge, Nishikatsura-mura, Mina-
mitsuru-gun (^tPlilPM#^H7 llf^); Ya-
manashi, HONSHU; 35°31'N, 138°49'E; re-
ported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 49:
XXXIII-30). See Fig. 2A.
Miuchisanrin, Ishii-mura, Sayo-gun (fcffilP^
^:|4^/il*mj^); Hyogo, HONSHU; 35°05'N,
134°23'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 30: XVIII-4). See Fig. 2A.
Miurayama vicinity, Outaki-mura, Nishichi-
kuma-gun O^S^PiJt^ftHMUjffe); Na-
gano, HONSHU; 35°48'N, 137°32'E; reported
in questionnaire survey conducted in 1923 by
K. Hasebe (Iwano, 1974, p. 45: XXXI-8). See
Fig. 2A.
Miyagi CM^), HONSHU; 37°47'-39°00'N,
140°16'-141°4rE; studied 1982-2002 at
various localities by Miyagi Monkey Investi-
gation Group (2003, p. 37). See Fig. 2B.
Miyajima-cho, Saeki-gun (fe-fSlP^SBlT); Hi-
roshima, MIYAJIMA; 34°16'N, 132°20'E;
population translocated from Shodoshima,
Kagawa Prefecture, in 1962; provisioned on
Miyajima (Iwamoto & Hirai, 1970, p. 395;
Shidei et al., 1981, p. 18). Birth season,
1962-1966, reported by Kawai et al. (1967, p.
38). Collected before 2002 by PRIKU staff;
PRJKU, 1 (skeleton only, with fragmented
skull). SW65.
Miyama National Forest, Ookawachi-mura, li-
nan-gun (ISl^?PAyRlrt^;^UjSWW); Mie,
HONSHU; 34°31'N, 136°28'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 37: XXIII-25). See
Fig. 2A.
Miyamoto, Takamori-machi, Shimoina-gun (T
PMW>mMViJ'B^y, Nagano, HONSHU;
35°34'N, 137°51'E; collected 6 Apr. and 14
May 1998 by M. Aimi; PRIKU, 2 (skeletons
only). NE76.
Miyamura, Takayama-mura, Kamitakai-gun (_h
it^lPSlil^EW); Nagano, HONSHU;
36°41'N, 138°24'E; collected 7 Nov. 1997-2
Nov. 1998 by M. Aimi; 4, (skeletons only).
Collected 19 Oct. 1998 by PRIKU staff;
PRIKU, 1 (skeleton only). NE49.
Miyanoura, Kamiyaku-cho, Kumage-gun (^^
W ± M X Vil 'B (D M ); Kagoshima
YAKUSHIMA 30°25'N, 130°34'E; collected 3
Dec. 1977 by PRIKU staff; PRIKU, 1 (skele-
ton only). SWIOO.
Miyanoura-Ambo. See Yakushima, Census Area
J.
Miyashiro, Hodaka-machi, Minamiazumi-gun
(^$*lP^SfflT^^); Nagano, HONSHU;
36°23'N, 137°49'E; collected 24 Aug. 1998 by
M. Aimi; PRIKU, 6 (skeletons only, including
2 with fragmented skulls). NE92.
Miyashita, Matsukawa-mura, Kitaazumi-gun (db
^•iP^AMIIW^T); Nagano, HONSHU;
35°34'N, 137°51'E; collected 22 Feb. 1998 by
PRIKU staff; PRIKU, 1 (skeleton only). NE76.
Miye. See Mie.
Mizuho-cho, Funai-gun (^n^lP^^fflJ); Kyoto,
HONSHU; ca. 35°10'N, 135°21'E; captured 26
Mar. 1977 by JMC staff, accidental death in
captivity 4 Apr. 1977; JMC, 1 (skull only).
SW39.
Mizunashiyama vicinity, Toga-mura, Higashito-
nami-gun (mfll):^lP^JM^7KMUjflfe); To-
yama, HONSHU; 36°17'N, 137°00'E; reported
in questionnaire survey conducted in 1923 by
K. Hasebe (Iwano, 1974, p. 42: XXIX-2). See
Fig. 2A.
Mizunoshitayama, Iseki, Nachi-mura, Higashi-
muro-gun (m#«^i9mW#Pl7Ky TUJ);
Wakayama, HONSHU; 33°39'N, 135°57'E;
reported in questionnaire survey conducted in
154
FIELDIANA: ZOOLOGY
1923 by K. Hasebe (Iwano, 1974, p. 34:
XX-45). See Fig. 2A.
Mobara-city ( j5S i^ TtT ); Chiba, HONSHU;
35°25'N, 140°18'E; Jomon subfossils (age,
2.3-12 Ka) reported by K. Kawanaka (1973, p.
\\5).NE129.
Mochihirasan vicinity, Futagawa-mura, Nishi-
muro-gun (ffi^ftlP-Jll^^^ajffe); Wa-
kayama, HONSHU; 33°52'N, 135°22'E; re-
ported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 33:
XX-17). SeeFig.2A.
Momobayama, Itoshiromi, Wakasa-cho,
Yazu-gun (A^^P^^fflJ.^ 6 B^^yXOj);
Tottori, HONSHU; 35°19'N, 134°25'E; re-
ported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 29:
XVII-2). See Fig. 2A.
Momozaki vcinity, Isato-mura, Minamimuro-gun
im^mW^m^mm^); Mle, HONSHU;
34°00'N, 136°03'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 37: XXIII-51). See Fig. 2A.
Moniwa-mura, Date-gun ('^ji^^j^SW); Fu-
kushima, HONSHU; 37°55'N, 140°27'E; re-
ported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 52:
XLI-10). SeeFig. 2A.
Morimatsu vicinity, Kitawauchi-mura, Minami-
muro-gun (]^#ft?Pdblfflrt^^^A^fte); Me,
HONSHU; 34°00'N, 136°14'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 37: XXIII-48). See
Fig. 2A.
Motoi vicinity, Higashiiyayama-son, Mima-gun
(H^liPmrn^Oj^TU^fte); Tokushima,
SHIKOKU; 33°54'N, 133°57'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 23: XI-22). See Fig.
2A.
Motokiyosumiyama. See Kiyosumiyama.
Motomura, Minamishinano-mura, Shimoina-gun
( T ^ 3P ?P 1^ if )I ^^ ^ ^ ); Nagano,
HONSHU; 35°18'N, 137°55'E; collected 20
Sep. 1998 by M. Aimi; PRIKU, 1 (skeleton
only). NE68. _
Motoyama, Muroto-cho, Aki-gun ($S^^P
ffll Tt UJ ); Kochi, SHIKOKU; 33°18'N,
134°10'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 22: X-44). See Fig. 2A.
Mt. Kiyosumi. See Kiyosumiyama.
Mt. Kuniwari. See Kuniwaridake; Nina- A.
Mt. Takago. See Takagoyama.
Mugikomoriyama vicinity, Ayukawa-mura, Ni-
shimuro-gun (M#«IPIiJIIW^3^^ajffe);
Wakayama, HONSHU; 33°41'N, 135°30'E;
reported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 33:
XX-22). See Fig. 2A.
Mugio, Yaku-cho, Kumage-gun (^^^MA.fflI
U^y, Kagoshima, YAKUSHIMA; 30°16'N,
130°36'E; collected 21 Jul. 1987 by PRIKU
staff; 3 (skeletons only). Collected 21-25 Jul.
1987 by PRIKU staff; PRIKU, 7 (postcranials
only). SW102.
Mukabakiyama, Minamikata-son, Higa-
shiusuki-gun {1^B^^ny5W<Jm\U); Mi-
yazaki, KYUSHU; 32°37'N, 131°34'E; re-
ported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 15: 11-25).
See Fig. 2A.
Mukaiyama, Mitsumata-mura, Mi-
namiuonuma-gun ( j^ ^ /S ?P H -(M W fS] UJ );
Niigata, HONSHU; 36°52'N, 138°47'E; re-
ported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 47:
XXXII-3). See Fig. 2A.
Mukawa-mura, Kitakoma-gun ( db ^ S IP ^ J 1 1
^ ); Yamanashi, HONSHU; ca. 35°45'N,
138°24'E; mtDNA samples collected before
1999 by Y. Kawamoto (1998, p. 54; 2002, p.
60). NE96.
Mumeinoyama, Imaichi-mura, Oono-gun (Ai?
lP4■T!T;f^t^. =S(DUj); Oita, KYUSHU;
33°09'N, 131°25'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 16: III-7). See Fig. 2A.
Mumeinoyama, Kawanobori-mura, Oono-gun
(AiflPJH^W^^CDUj); Oita, KYUSHU;
33°00'N, 131°44'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 16: 1II-8). See Fig. 2A.
Mumeinoyama, Onoichi-mura, Oono-gun (Ai?
W'\^m'i^^M'^<D\l\y, Oita, KYUSHU;
32°5rN, 131°36'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 16: III-l). See Fig. 2A.
FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES
155
Mumeinoyama. Shigeoka-mura, Oono-gun (AS
^M^^M4 (D iU); Oita, KYUSHU;
32°48'N, 131°36'E; reported in questionnaire
sur\ey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 16: III-2). See Fig. 2A.
Mureyama \icinit\'. Wabuka-mura, Nishi-
muro-gun (ffi^ilP^}^^#^Lajfte); Wa-
kmama. HONSHU; 33°3rN, 135°40'E; re-
ported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano. 1974, p. 34:
XX-34). See Fig. 2A.
Muroji. Muro-mura, Uda-gun (^PSIP^^^^
^#); .Vara, HONSHU; 34°32'N. 136°02'E;
blood samples collected before 1992 by No-
zawa et al. (1991. p. 414; 1996, p. 6). SW21.
Murokawayama. Tsuki-mura, Arida-gun (WBQIP
^ ^ tf ^ Jll Ol ); ^Fa^mw/;a, HONSHU;
33°58'N. 135°14'E; reported in questionnaire
sur\ey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 33: XX-6). See Fig. 2 A.
Myogisan National Forest. Mvogi-machi. Kita-
kaJu-a-gun {it^^m^'m^i^mium^^)-
Gunma, HONSHU; 36°16'N. 138°47'E: re-
ported in questionnaire sur\ey conducted in
1923 by K. Hasebe (Iwano. 1974. p. 51: XL-4).
See Fig. 2A.
Myojinyama. Mitono-mura, Kitashitara-gun (it
^^Wmm^mnOi); Aichi. HONSHU;
35°05'N. 137°40'E; reported in questionnaire
sur\ey conducted in 1923 by K. Hasebe
(Iwano. 1974. p. 40: XXVII-5). See Fig. 2A.
Myomisan. Youka-mura, Yabu-gun (^5<^1PA
M,^^^\h)\ Hyogo, HONSHU; 35°24'N,
134°45'E; reported in questionnaire survey
conducted in 1 923 by K. Hasebe (Iw ano. 1 974,
p. 30: XVIII- 17). See Fig. 2A.
Nabegatani. Chikusa-mura, Shisou-gun (t^^IP
=fU^t^r^): Hyogo, HONSHU; 35°08'N.
134°28'E: reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 30: XVIII-7). See Fig. 2A.
Nabemashi vicinity. Shirahagi-mura, Nakanii-
kawa-gun ( 4^ ff J 1 1 IP 6 M^^i^^): Toyama,
HONSHU; 36°39'N, 137°30'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano. 1974, p. 42: XXIX-6). See Fig.
2A.
Nadanishiyama \ icinity. Higashimata-mura, Ta-
kaoka-gun (iS[55iPm3^^1JSajfte); Kochi,
SHIKOKU; 33° 1 3?^, 133°14'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 22: X-17). See Fig.
2A.
Nadayama, Tsubaki-son, Naka-gun (SPMIP^^
M\h): Tokushima. SHIKOKU; 33°49'N,
134°40'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974.
p. 23: XI- 12). See Fig. 2A.
Nadayama vicinity, Kaminokae-cho, Taka-
oka-gun (il[5^^±y ia>lffllliajfte); Kochi,
SHIKOKU: 33°16'N, 133°16'E: reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 22: X-18). See Fig.
2A.
Nagaflika vicinity, Handairayama-son,
Mima-gun (ll.l?P¥¥UJ^^)^^); Toku-
shima, SHIKOKU; 33°58'N, 134°08'E; re-
ported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 23: XI-32;
Mito, 1989, p. 25). See Fig. 2A.
Nagahashiri vicinity. Hideya-mura, Higashikan-
bara-gun (m^i^lP B tli^^^^'^fe); Nii-
gata, HONSHU; 37°43'N, 139°32'E; reported
in questiormaire survey conducted in 1923 by
K. Hasebe (Iwano, 1974, p. 47: XXXII- 19).
See Fig. 2A.
Nagakura vicinity, Hase-mura, Kanzaki-gun (tt
^^^"^m-llO^^Y Hyogo, HONSHU;
35°07'N, 134°44'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974. p. 30: XVIII- 14). See Fig. 2A.
Nagano (MS) (= Shinshiu), HONSHU;
35°12'-37°01'N, 137°13'-138°48'E; collected
Sep. 1882 by P. L. Jouy; USNM, 1 (skull only).
Questionnaire survey conducted June 1972 by
E. Tokita and S. Hara (1977, p. 27). Not
mapped.
Nagaoi. Egawasaki-mura. Hata-gun (♦S^^PxIl
jl|*f^k^); Kochi, SHIKOKU; 33°irN,
132°47'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 21: X- 13). See Fig. 2 A.
Nagasaka-mukaiyama. Takigawa-mura,
Naga-gun {4%^M}\\-^^^^\Uy, Mie,
HONSHU; 34°34'N, 136°05'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 36: XXIII- 17). See
Fig. 2A.
156
FIELD! AN A: ZOOLOGY
Nagata, Kamiyaku-cho, Kumage-gun (^^?P_h
MrXfflJ^ffl); Kagoshima, YAKUSHIMA; ca.
30°23'N, 130°25'E; collected 7 Jul. 1977 and
22 Jul. 1987 by PRIKU staff; PRIKU, 2 (1
skeleton without mandible, 1 postcranials
only). Collected 20 Jul. 1987 by Mr. Imaegawa;
PRIKU, 2 (skeletons only). Collected 25 Jul.
1987 by Mr. Sinmachi; PRIKU, 1 (skeleton
only). Collected 25 Feb. 1989 by PRIKU staff;
PRIKU, 4 (skeletons only). SWIOI.
Nagata-Issou. See Yakushima, Census Area 1 .
Nagata-todai, Kamiyaku-cho, Kumage-gun (^
^?P±Jl:^fflT*Baff^); Kagoshima,
YAKUSHIMA; ca. 30°23'N, 130°23'E; ob-
served Aug. 1978 by T. Maruhashi (1982, p.
318; cf. Takasaki & Masui, 1984, p. 311).
SWlOl.
Nagato-okuyama, Fukusawa-mura, Kaminii-
kawa-gun (±ifJII^II)K^m^f MUJ); To-
yama, HONSHU; 36°25'N, 137°19'E; reported
in questionnaire survey conducted in 1923 by
K. Hasebe (Iwano, 1974, p. 42: XXIX-3). See
Fig. 2A.
Nagatokyo vicinity, Ikumo-son, Abu-gun (PrI^
^ ^m^ ^f^^ - "^ )\ Yamaguchi,
HONSHU; 34°23'N, 131°37'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 25: XIII-8). See Fig.
2A.
Nagatokyo vicinity, Kawakami-son, Abu-gun
{n^W>\\\:t^^f^^-w)\ Yamaguchi,
HONSHU; 34°21'N, 131°28'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 25: XIII-7). See Fig.
2A.
Nagatokyo vicinity, Shinobu-son, Abu-gun (Pr[
^W^U^^^cf^m-^y, Yamaguchi,
HONSHU; 34°20'N, 131°36'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 25: XIII-9). See Fig.
2A.
Nagatoro. See Houtosan.
Nagisoyama, Azuma-mura, Nishichikuma-gun
(SmiSlPl^g^^TK^Uj); Nagano,
HONSHU; 35°35'N, 137°38'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 45: XXXI- 1). See Fig.
2A.
Nagis(wama vicinity, Yomikaki-mura, Nishichi-
kuma-gun (S^SiPlTcH^lt^^ajfe);
Nagano, HONSHU; 35°37'N, 137°38'E; re-
ported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 45:
XXXI-2). See Fig. 2A.
Nahisanrin, Aioi-mura, Gujo-gun (?P_hlP1S^
^ SP it UJ ^ ); Gifu, HONSHU; 35°42'N,
136°56'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p.41:XXVIII-23). SeeFig. 2A.
Naka, Natashou-mura, Onyu-gun (]ai![?P^B9
Ji^ct'); Fukui, HONSHU; 35°23'N, 135°38'E;
collected Aug.-Dec. 1994 by PRIKU staff;
PRIKU, 3 (skeletons only). SW42.
Nakabusa National Forest, Ariake-mura, Mi-
namiazumi-gun (]^$#IPWB^^4'MSW
^); Nagano, HONSHU; 36°21'N, 137°50'E;
reported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 46:
XXXI-17). SeeFig.2A.
Nakadachi, Minamishinano-mura, Shimoina-gun
( T # IP ?P ^ € ^M ^ 4^ it ); Nagano,
HONSHU; 35°22'N, 137°58'E; collected 9 Sep.
1998 by M. Aimi; PRIKU, 1 (skeleton only).
NE69.
Nakahama vicinity, Shimizu-mura, Hata-gun (Wl
^?P)t7K;t4 4'^^); Kochi, SHIKOKU;
32°46'N, 132°58'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 21: X-6). See Fig. 2 A.
Nakahora, Ooshika-mura, Shimoina-gun (T#SP
W>XU^^M); Nagano, HONSHU; 35°33'N,
138°02'E; collected 12 Apr. 1998 by M. Aimi;
PRIKU, 1 (skeleton only). NE70.
Nakakimura, Usui-machi, Usui-gun (fift:^?PS
#fflI4^^^); Gunma, HONSHU; 36°18'N,
138044'E; reported in questionnaire survey
conducted in 1 923 by K. Hasebe (Iwano, 1 974,
p. 51:XL-5). SeeFig. 2A.
Nakakodomariyama, Kodomari-mura, Ki-
tatsugaru-gun ( db ^ ^ ?P 'J^ )& ^ 4^ ^^ )^ Ol );
Aomori, HONSHU; 41°07'N, 140°20'E; re-
ported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 57:
XLVI-12). SeeFig. 2A.
Nakakurayama, Shimoyakawa-mura, Aga-
wagun ( f^ Jll iP T A Jll^ 4^^111); Kochi,
SHIKOKU; 33°36'N, 133°20'E; reported in
questionnaire survey conducted in 1923 by K.
FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES
157
Hasebe (Iwano, 1974, p. 22: X-24). See Fig.
2A.
Nakama, Yaku-cho, Kumage-gun (^^?PM^
BTTft^fB^); Kagoshima, YAKUSHIMA; 30°16'N,
130°29'E; collected 20 Jul. 1987 by Mr.
Imaegawa; PRIKU, 1 (skeleton only). Col-
lected 26-27 Jul. 1987 by PRIKU staff;
PRIKU, 3 (postcranials only). Collected 26-27
Jul. 1987 by Mr.Yasumaki; PRIKU, 7 (3
skeletons only, 4 skulls only). Collected 27 Jul.
1987 by PRIKU staff; PRIKU, 1 (skeleton
only). Collected 27 Jul. 1987 by Mr. Mura-
kami; PRIKU, 1 (skull only). SW102.
Nakamura-city (4^^TtJ); Kochi, SHIKOKU; ca.
32°59'N, 132°56'E; mtDNA samples collected
before 2003 by Y. Kawamoto (2002, p. 60).
SW79.
Nakanogoudani vicinity, Samoto-mura, Nishi-
muro-gun (ffi#«lPfe^^^f >f ^ ^P#ffe);
Wakayama, HONSHU; 33°35'N, 135°38'E;
reported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 34:
XX-31). SeeFig. 2A.
Nakao, Tomisato-mura, Uma-gun (^JSIPS^P
W 4^ M ); Ehime, SHIKOKU; 33°53'N,
133°3rE; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 18:IX-10). SeeFig. 2A.
Nakaodani vicinity, Go-mura, Arida-gun (Wffl
W^^^W^^y, Wakayama, HONSHU;
34°03'N, 135°25'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 33: XX-5). See Fig. 2A.
Nakase, Yaku-cho, Kumage-gun (^^^M^fflJ
4^^); Kagoshima, YAKUSHIMA; 30°16'N,
130°36'E; collected 23 Jul. 1987 by PRIKU
staff; PRIKU, 3 (2 skeletons only, 1 postcrani-
als only). SW102.
Nakasegawa, Yaku-cho, Kumage-gun (^^^M
X^^M]\\); Kagoshima, YAKUSHIMA;
30°16'N, 130°36'E; collected 21 Jul. 1987 by
PRIKU staff: PRIKU, 1 (skeleton only).
SW102.
Nakasu, Suwa-city ( M IS rfT 4^ /jtl ); Nagano,
HONSHU; 35°59'N, 138°07'E; collected 26
Jul. 1998 by PRIKU staff; PRIKU, 1 (man-
dibular fragments and postcranials only).
NE95.
Nakatsugawa vicinity, Taisho-mura, Hata-gun
(M^aPAlE^^4^}tJI|'te); Kochi, SHIKOKU;
33°14'N, 132°59'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 21: X-15). See Fig. 2A.
Nakatsugawayama, Sakanoshita-mura, Su-
zuka-gun (Ip El^^feT W 4^ /$)rI Uj ); Mie,
HONSHU; 34°53'N, 136°23'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 36: XXIII- 10). See
Fig. 2A.
Nakayama, Takayama-mura, Kamitakai-gun (_h
K#lPI^Uj^4^LiJ); Nagano, HONSHU;
36°4rN, 138°23'E; collected 19 Oct. 1998 by
M. Aimi, PRIKU, 1 (skeleton only). Collected
Nov. 1998 by PRIKU staff; PRIKU, 1 (skele-
ton only). NE49.
Namesawayama, Mitomi-mura, Higashiyamana-
shi-gun (^UJ^^PHSW^/RUJ); Yamanashi,
HONSHU; 35°51'N, 138°45'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 48: XXXIII-20). See
Fig. 2A.
Nametoko, Nogawa, Uwajima-city (^^PfiTtTi?
JIlTiJ^); Ehime, SHIKOKU; ca. 33°12'N,
132°38'E; provisioned group; birth season,
1963-1966, reported by Kawai et al. (1967, pp.
37, 39). Blood samples collected before 1992
by Nozawa et al. (1991, p. 414; 1996, p. 6).
External measurements taken by Hamada et al.
(1996a, pp. 98, 99). MtDNA samples collected
before 2003 by Y. Kawamoto (2002, p. 60).
SIV77.
Namisegawara, Kitaura-son, Higashiusuki-gun
{MB^Witm^^&mniW^y, MiyazaU,
KYUSHU; coordinates unknown; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Mito,1989, p. 25). Not mapped.
Nangai-mura, Senboku-gun (-(llldb^P I^^KW);
Akita, HONSHU; 39°24'N, 140°24'E; mtDNA
extracted from umayazaru (talismanic hunting
trophy) before 2003 by T. Agatsuma and M.
Ishigami (2002, p. 79). NEI5 .
Nansei-cho, Watarai-gun (^^1^1^^ BJ); Mie,
HONSHU; ca. 34°20'N, 136°41'E; observed
Aug. 1979 by K. Masui (Takasaki & Masui,
1984, pp. 310, 317). 5^2i.
Naruse. See Narusegawa.
Narusegawa, ( 1l 0 Jl| ); Miyagi, HONSHU;
38°30'N, 141°04'E; observed Jul. 2002 by K.
158
FIELDIANA: ZOOLOGY
Izawa, T. Uno, and H. Fujita (2003, p. 27).
NE23.
Natasho-mura, Onyu-gun (3si!(?P=SfflJiW);
Fukiii, HONSHU; ca. 35°24'N, 135°41'E;
mtDNA samples collected before 2003 by Y.
Kawamoto (2002, p. 60). SW42.
National Forest, Akaishi-mura, Nishitsugaru-gun
(ffi)$giP#;5;|^S#;i9^); Aomori, HONSHU;
40°44'N, ]40°irE; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 57: XLVI-6). See Fig. 2 A.
National Forest, Fukaura-mura, Nishitsugaru-gun
0)*fe?P;^)t^^aW^); Aomori, HONSHU;
40°37'N, 139°56'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 57: XLVI-2). See Fig. 2A.
National Forest, Hokujo-mura, Kitaazumi-gun
( Jb S « ^ db ^ ^ S W ^ ); Nagano,
HONSHU; 36°40'N, 137°50'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 46: XXXI-22). See
Fig. 2A.
National Forest, Imabetsu-mura, Higashit-
sugaru-gun (^^/^^^^SIJ^SW^); Aomori,
HONSHU; 41°10'N, 140°29'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 57: XLVI-14). See
Fig. 2A.
National Forest, Iwasaki-mura, Nishitsugaru-gun
(M}$^IP^llit^SIW^); Aomori, HONSHU;
40°34'N, 139°57'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 57: XLVI-1). See Fig. 2A.
National Forest, Kamishiro-mura, Kitaazumi-gun
( db ^ » ?P tt J^ ^^r il W ^ ); Nagano,
HONSHU; 36°38'N, 137°49'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 46: XXXI-21). See
Fig. 2A.
National Forest, Matsukawa-mura, Kitaa-
zumi-gun (db^ft^P^iJII^SW^I^^); Nagano,
HONSHU; 36°25'N, 137°50'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 46: XXXI- 18). See
Fig. 2A.
National Forest, Minamiaiki-mura, Mina-
misaku-gun (l^fe^^Pi^^lTK^ttH^^);
Nagano, HONSHU; 36°00'N, 138°35'E; re-
ported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 46:
XXXI-37). See Fig. 2A.
National Forest, Nakatsuchi-mura, Kitaa-
zumi-gun (db$#?P'^±^g#^); Nagano,
HONSHU; 36°50'N, 137°57'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 46: XXXI-23). See
Fig. 2A.
National Forest, Nishishiwa-son, Kamo-gun (I?
j^?PIS.^^^^S##); Hiroshima, HONSHU;
34°29'N, 132°36'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 26: XIV- 16). See Fig. 2A.
National Forest, Oodose-mura, Nishitsugaru-gun
(S/$^1PAF^^SW#); Aomori,
HONSHU; 40°43'N, 140°04'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 57: XLVI-5). See Fig.
2A.
National Forest, Taira-mura, Kitaazumi-gun (db
$#1P¥^1#^); Nagano, HONSHU;
36°33'N, 137°49'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 46: XXXI-20). See Fig. 2 A.
National Forest, Tairadate-mura, Higashit-
sugaru-gun (K/$^1P¥^^@'^#); Aomori,
HONSHU; 41°07'N, 140°37'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 57: XLVI-1 5). See
Fig. 2A.
National Forest, Tokiwa-mura, Kitaazumi-gun
(db^ftlPSM^lW*^); Nagano,
HONSHU; 36°28'N, 137°49'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 46: XXXI-19). See
Fig. 2A.
National Forest, Yuzono, Suki-son, Nishimoro-
kata-gun (IS^miP^^Wtt^SW^); Mi-
yazaki, KYUSHU; 32°05'N, 131°05'E; re-
ported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 15: 11-14).
See Fig. 2A.
National Forest vicinity, Kameyama-mura, Ki-
mitsu-gun (^)*lPmiljWSW^^); Chiba,
HONSHU; 35°13'N, 140°07'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 50: XXXVI-4). See
Fig. 2A.
Natsusaka vicinity, Kitayama-mura, Yama-
FOODEN AND AIM!: SYSTEMATIC REVIEW OF JAPANESE MACAQUES
159
gata-gun (OjmiPdbOJ^MJS^); Gifu,
HONSHU: 35°40'N, 136°47'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 41: XXVIlI-14). See
Fig. 2A.
Negitani. Nishikawa-mura, Gujo-gun (?P_hlPS
]\\ ^ :^ =^' ^ ): Gifu, HONSHU: 35°48'N.
136°53'E: reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano. 1974.
p. 41 : XXVIII-25). See Fig. 2 A.
Nenbutsudani National Forest, Minochi-son.
Saeki-gun {^\^^:^P^^±\U"^mM^Y
Hiroshima. HONSHU: 34°31'N. 132°18'E:
reported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano. 1974. p. 26:
XIV-2). See Fig. 2A.
Neo-mura, Motosu-gun (^^IP^^^): Gifu.
HONSHU: ca. 35°38'N, 136°37'E: mtDNA
samples collected before 2003 by Y. Kawa-
moto (2002, p. 60). NE59.
Nezumiana, Matsukawa-mura. Kitaazumi-gun
(db^^^P^A^JII^ETv:): Nagano, HONSHU:
36°24'N, 137°50'E: collected 22 Feb. 1998 by
M. Aimi: PRIKU, 2 (skeletons only). NE92.
Nigure, Kamianama-mura. Oono-gun (ASlP-h
7^,l^if •): Fukui, HONSHU: 35°51'N.
136°46'E: reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano. 1974.
p. 39: XXV-22). See Fig. 2A.
Nikko-city (B^fellT): Tochigi. HONSHU: ca.
36°45'N, 139°37'E: albino monkey capuired
before 1826 (von Siebold, MS., 1830-1842, p.
1: cf. Temminck. 1842. p. 10): specimen not
preserved. Collected in 1875 by Mr. v.
Hilgendorf (Schweyer, 1909. p. 8): ZMB. 8 (1
skeleton only. 7 skulls only). Eight skulls pur-
chased in 1900-1901 by K. A. Haberer
(Schweyer. 1909. p. 8): some or ail of these
skulls may be included among unlocalized
specimens examined in AIUM. Reported pre-
sent before 1939 by N. Kuroda (1938. p. 112:
cf. Study Group on the Present Status of Japa-
nese Monkeys, 1977b, p. 26). Collected 8 Sep.
1981-24 Sep. 1986 by TPM staff: TPM. 21
(skulls only, excluding 10 specimens not seen,
data from specimen list). MtDNA samples
collected ca. 1986-1991 by Hayasaka et al.
(1991, p. 400). Blood samples collected before
1992 by Nozawa et al. (1991, p. 414; 1996, p.
6). External measurements taken by Hamada et
al. (1996a, pp. 98, 99). MtDNA samples col-
lected before 1998 by Y. Kawamoto (1997, p.
33:2002.p. 60).A'£-^i.
Nikkosan. Yahata-mura. Arida-gun (WB9?PANI
^S3feaj): Wakayawa, HONSHU: 34°05'N,
135°27'E: reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano. 1974,
p. 33:XX-4). SeeFig. 2A.
Nina-A. Hanyama. Kamiyaku-cho. Kumage-gun
(^^lP±M:XfflI¥aj~:)-A||): Kagoshiiiw,
YAKUSHIMA: 30°21'N. 130°23'E: field study
conducted Aug.-Dec. 1976 by T. Maruhashi
(1981, p. 2: cf.^982, p. 323; Soltis et al., 2000,
p. 196). Feeding association with Cenus nip-
pon observed Jan.-May 2002 by B. Majolo
and R. Ventura (2004. p. 35). SWIOI.
Ninomiya-machi. Naka-gun ( "^ ?P — §' fflj ):
Kanaga^va, HONSHU: ca. 35°18'N, 139°16'E;
mtDNA samples collected before 1998 by Y.
Kawamoto (1997. p. 33). NE105.
Nireyama vicinity. Nire-mura, Kamitakai-gun
( ± it ^ IP t ^L ^ t ^L OJ fife ): Nagano.
HONSHU: 36°36'N. 138°21'E: reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano. 1974. p. 46: XXXI-31). See
Fig. 2A.
Nisakasanrin vicinity. Nigou-mura, Kita-
muro-gun {i\L^UW-n^n^\^^^)\ Mie.
HONSHU: 34°13'N. 136°20'E: reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974. p. 37: XXIII-40). See
Fig. 2A.
Nishi. Ooshika-mura, Shimoina-gun (T'^SP^P
AE^ffi): Nagano. HONSHU: 35°35'N,
138°02'E: collected 4 Dec. 1998 by PRIKU
staff: PRIKU. 1 (skeleton only). NE70.
Nishiazai-cho. Ika-gun (##?PS^#fflI):
Shiga. HONSHU: ca. 35°29'N. 136°08'E:
mtDNA samples collected before 2003 by Y.
Kawamoto (2002. p. 60). SW4.
Nishifijjiyama. Kami-mura, Shimoina-gun (T*^
IP ?P ± :14 S S Oj ): Nagano. HONSHU;
35°24'N, 137°58'E: reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974. p. 47: XXXI-47). See Fig. 2A.
Nishiharuchika, Ina-city ("^IP rf5"ffi#]fi); Na-
gano, HONSHU; 35°47'N, 137°56'E; collected
17 Mar.-I3 Aug. 1998 by M. Aimi; PRIKU,
160
FELDIANA: ZOOLOGY
1 1 (skeletons only, including 4 with frag-
mented skulls). NE79.
Nishihikasa, Kimitsu-city ( ^ /$ TtT g 0 ^ );
Chiba, HONSHU; 35°13'N, 139°59'E; col-
lected 22 Aug. 1998 by PRIKU staff; PRIKU,
1 (skeleton only). NE120.
Nishihinobori, Kisuki-cho, Ohara-gun ( AJ^^^
;^fflIMS^); Shimane, HONSHU; 35°15'N,
132°54'E; collected. 4 Dec. 1994 and 12 Feb.
1995 by PRIKU staff; PRIKU, 2 (skeletons
only). SW59.
Nishiizu-cho, Kamo-gun (Mj^^PM^aBI);
Shizuoka, HONSHU; 34°46'N, 138°47'E;
mtDNA samples collected before 2003 by Y.
Kawamoto (2002, p. 60). NEIOO.
Nishikatsura-cho, Minamitsuru-gun (j^li^li^
S^fflJ); Yamanashi, HONSHU; ca. 35°30'N,
138°5rE; mtDNA samples collected before
2003 by Y. Kawamoto (2002, p. 60). NE98.
Nishikomagadake, Ina-machi, Kamiina-gun (_b
#ilP?P^ilWM|6)^l&); Nagano, HONSHU;
35°50'N, 137°56'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 46: XXXI-41). See Fig. 2A.
Nishikurokiyama. See Higashi- and Ni-
shi-Kurokiyama.
Nishimeya-mura, Nakatsugaru-gun (^'/♦S^PM
iM^); Aomori, HONSHU; ca. 40°30'N,
140°15'E; captured alive, briefly held in cap-
tivity, donated to JMC 22 Jul. 1966; JMC, 1
(skull only). NEW.
Nishine, Kitaokuni-mura, Nishiokitama-gun (ffi
amPdb^hS^ffi^); Yamagata, HONSHU;
38°10'N, 139°46'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 55: XLII-4). See Fig. 2A.
Nishinomaki, Saimoku-mura, Kitakanra-gun (db
^^^'^'^^'Em^y, Gimma, HONSHU;
36°14'N, 138°40'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 51: XL-3). See Fig. 2A.
Nishinoyakaeyama, Saigou-son, Higa-
shiusuki-gun {1^B^WmM¥^'^ )\^\\^);
Miyazaki, KYUSHU; 32°24'N, 131°28'E; re-
ported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 15: 11-21).
See Fig. 2A.
Nishiogawa vicinity, Uchitomi-mura, Onyu-gun
( ':&m. ?P 1*1 ^Wi ^ H d^ Jll fife ); Eukui,
HONSHU; 35°31'N, 135°47'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 39: XXV- 1 1). See Fig.
2A.
Nishiohira, Ichibu-mura, Kuma-gun (^^IP —
E^ffiA^); Kumamoto, KYUSHU; 32°10'N,
130°52'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 16:IV-2). SeeFig. 2A.
Nishitokoo vicinity, Murohani-mura, Izushi-gun
(tbSlP^il^S^Mffe); Hyogo, HONSHU;
35°26'N, 134°5rE; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 30: XVIII- 18). See Fig. 2 A.
Nishitoyonaga-mura, Nagaoka-gun (:M[5??PMM
^W); Kochi, SHIKOKU; 33°49'N, 133°45'E;
reported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 22: X-34).
See Fig. 2A.
Nishiyama, Minamiwada-mura, Shimoina-gun
(T^jiPiP^^ffl^ffiOj); Nagano,
HONSHU; 35°19'N, 137°55'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 47: XXXI-50). See
Fig. 2A.
Nishiyama, Wada-mura, Shimoina-gun (T#3P
iP^ffl^ffiOj); Nagano, HONSHU; 35°19'N,
137°57'E; reported in questionnaire survey
conducted in 1 923 by K. Hasebe (Iwano, 1 974,
p. 47: XXXI-49). See Fig. 2A.
Niu-mura, Ika-gun (##IP:I^^^); Shiga,
HONSHU; 35°34'N, 136°15'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 38: XXIV- 12). See
Fig. 2A.
Nonesankei, Sakihama-mura, Aki-gun ($S?P
feSMWS'aaj.^); Kochi, SHIKOKU;
33°28'N, 134°09'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 22: X-43). See Fig. 2A.
Nonoboriyama vicinity, Nonobori-mura, Su-
zuka-gun ($^SiPi?^^i?^aJ^); Mie,
HONSHU; 34°56'N, 136°25'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 36: XXIII-9). See Fig.
2A.
Nonogawa vicinity, Nishikamiyama-mura,
Hata-gun (^i^iPS±aj:^i?<? JHffe); Kochi,
SHIKOKU; coordinates unknown; reported in
FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES
161
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 22: X-15). Not
mapped.
Norogawairi, Ashiyasu-mura, Nakakoma-gun
{^mmW>P^^m^)\\X); Yamanashi,
HONSHU; 35°40'N, 138°17'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 48: XXXIII-11). See
Fig. 2A.
Northern Japan Alps, Kitaazumi-gun and Mi-
namiazumi-gun {i\:T )l7°7s), 600-3000 m
elevation; Nagano, HONSHU; ca. 36°10-30'N,
137°40-50'E; 22 troops studied 26 Dec.
1996-21 Feb. 2000 by S. Izumiyama et al.
(2003, p. 465). Not mapped (see Tatai and
nearby localities).
Northern villages, Tsuno-gun (tPii^Pdb^^^);
Yamaguchi, HONSHU; 34°14'N, 131°51'E;
reported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 25:
XIII-12). SeeFig. 2A.
Nozokinosawa vicinity, Kasshi-mura, Kami-
hei-gun (±r^#lP¥^Wy V^y)Rffe);
Iwate, HONSHU; 39°14'N, 14r49'E; reported
in questionnaire survey conducted in 1923 by
K. Hasebe (Mito, 1989, p. 27). See Fig. 2A .
Nuidouishiyama, Sai-mura, Shimokita-gun (Tdb
fPfe^^i^jiSOj); Aomori, HONSHU;
41°23'N, 140°53'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 57: XLVI-22). See Fig. 2A.
Nutazawa, Kimitsu-city (:^/$rfT.^.ffl)H); Chiba,
HONSHU; 35°12'N, 140°01'E; collected 20
Aug. and 16 Dec. 1998 by PRIKU staff;
PRIKU, 2 (skeletons only). NE120.
Nyudougatake, Tsubaki-mura, Suzuka-gun (IpM
fP^^AM;!)'"*^); Me, HONSHU; 34°58'N,
136°26'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 36: XXIII-7). See Fig. 2A.
Nyudouzan vicinity, Toyohara-mura, Nishi-
muro-gun (ffi^SlPMJ^^Ajllijffe); Wa-
kayama, HONSHU; coordinates unknown; re-
ported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 34:
XX-38). Not mapped.
Obi, Masutomi-mura, Kitakoma-gun (Jb^S^
iis^^J^M); Yamanashi, HONSHU; 35°52'N,
138°3rE; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 48: XXXIlI-18). See Fig. 2A.
Ochiai, Ooshika-mura, Shimoina-gun (T'^SPIP
AEW^H-); Nagano, HONSHU; 35°34'N,
138°02'E; collected 14 Dec. 1998 by PRIKU
staff; PRIKU, 2 (skeletons only). NE70.
Odairayama vicinity, Hasegawa-mura, Oono-gun
(Ai??P^#JI|^m¥ajffe); Oita, KYUSHU;
32°52'N, 131°25'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 16: III-3). See Fig. 2A.
Odake, Kurihashi-mura, Kamihei-gun (_h^'(^?P
^M^^^); Iwate, HONSHU; 39°20'N,
141°43'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Mito, 1989,
p. 25). See Fig. 2A.
Odawara-city ( ^J"* ffl J^ rfT ); Kanagawa,
HONSHU; ca. 35°15'N, 139°10'E; mtDNA
samples collected before 1998 by Y. Kawa-
moto (1997, p. 33; 2002, p. 60). NEJ04.
Odo, Ohtsuki-cho, Hata-gun (<#^?PA>^ fflJA
^); Kochi, SHIKOKU; ca. 32°47'N, 132°38'E;
blood samples collected before 1992 by No-
zawa et al. (1991, p. 414; 1996, p. 6). SW80.
Odomisaki, Ohtsuki-cho, Hata-gun (^^^1PA>^
ffll A ^ llif ); Kochi, SHIKOKU; 32°46'N,
132°38'E; external measurements taken by
Hamada et al. (1996a, pp. 98, 99). SW80.
Ofunato-city (A^p^y^gTfT); Iwate, HONSHU; ca.
39°04'N, 14r43'E; mtDNA samples collected
before 1998 by Y. Kawamoto (1997, p. 33).
NE21.
Ogachi-machi, Ogachi-gun ( ^ 0 ?P 4i 0 fflj );
Akita, HONSHU; ca. 39°00'N, 140°19'E;
mtDNA samples collected before 1998 by Y.
Kawamoto (1997, p. 33; 2002, p. 60). NEI7.
Ogadake, Miyori-mura, Shioya-gun (^^IPH
i^^^^^y, Tochigi, HONSHU; 36°58'N,
139°4rE; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 51:XXXIX-6). SeeFig. 2A.
Ogawa-mura, Higashimuro-gun (^^ftlPd'^JH
^ ); Wakayama, HONSHU; 33°36'N,
135°48'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 34:XX-41). SeeFig.2A.
Ogawadani, Yamazaki-mura, Shimoniikawa-gun
( T ^ Jll ?P UJ ^ ^ ^h Jll # ); Toyama,
HONSHU; 36°55'N, 137°34'E; reported in
162
FIELDIANA: ZOOLOGY
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 42: XXIX- 11). See
Fig. 2A.
Ogawaoku vicinity, Kamikawa-mura, Minami-
muro-gun (^#■glP#JI|^^tmJllllfte); Mie,
HONSHU; 33°56'N, 136°00'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 37: XXIII-52). See
Fig. 2A.
Ogikubo, Takayama-mura, Kamitakai-gun (-hi§
^M>m\h¥i^^)\ Nagano, HONSHU;
36°41'N, 138°25'E; collected 26 Sep.-12 Nov.
1997 by M. Aimi; PRIKU, 2 (skeletons only).
Collected 7 Aug. 1998 by PRIKU staff;
PRIKU, 1 (skeleton only, with fragmented
skull). NE49.
Ohamidake ( ADtlS), 3050 m elevation; 36°20'N,
137°39'E; Nagano, HONSHU; reported pre-
sent before 1988 by S. Izumiyama (1987a, p. 8;
\9^1h,p.A2).NE88.
Ohara vicinity, Kitatani-mura, Oono-gun (Ai?
IPdk^Wd^Jiffe); Fukid, HONSHU; 36°05'N,
136°35'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 39: XXV-20). See Fig. 2A.
Oharadake, Yachi-mura, Ichishi-gun (— ^n^A
^ ^ d^ J^ H ); Mie, HONSHU; 34°33'N,
136°15'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 37:XXIII-19). SeeFig. 2A.
Ohira National Forest vicinity, Kobayashi-cho,
Nishimorokata-gun (SMli^^d^^fflTA^S
M ^^); Miyazaki, KYUSHU; 32°00'N,
131°00'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 15:11-13). SeeFig. 2A.
Ohirayama, Inuyama-city (i^lilTfiA^^lll); Aichi,
HONSHU; 35°23'N, 136°56'E; variably provi-
sioned population, translocated from Yaku-
shima in Mar. 1957 (Kawai, 1960, pp. 204,
222; cf. Fooden & Aimi, 2003, p. 111). Birth
season, 1957-1966, reported by Kawai et al.
(1967, p. 39). Blood samples collected before
1992 by Nozawa et al. (1991, p. 414; 1996, p.
7). NE6L
Ohsawakanrin, Hourai-mura, Kitakoma-gun (db
&0?Pm5lt^A)HtW); Yamanashi,
HONSHU; 35°50'N, 138°16'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 48: XXXIII- 16). See
Fig. 2A.
Oide, hills near. See Omachi, hills near.
Oitsuge vicinity, Nishiki-mura, Kitamuro-gun
(db^SiPII^7t->r7<rffe); Mie, HONSHU;
34°13'N, 136°25'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 37: XXIII-39). See Fig. 2A.
Okinoshima, Hazu-cho, Hazu-gun (fSs^Piffifi
fflT/4'(7)S); Aichi, OKINOSHIMA; 34°44'N,
137°10'E; provisioned population, translocated
from Shodoshima via JMC to Okinoshima in
May 1957; birth season, 1958-1966, reported
by Kawai et al. (1967, p. 38); translocated
from Okinoshima to JMC in Jan. 1998. NE62.
Okinoyama, Yamagata-son, Yazu-gun (AM?P
ajJf^^/4'yULl); Tottori, HONSHU; 35°16'N,
134°18'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 29:XVII-1). SeeFig. 2A.
Okoppe River, upper, Oma-machi, Shi-
mokita-gun (TdklP APeW^P Jll±)riit);
Aomori, HONSHU; ca. 41°26'N, 140°57'E;
reported 27-28 Jun. 1970 by Working Group
for the Conservation of Japanese Monkeys
(1970, p. 401; cf Study Group on the Present
Status of Japanese Monkeys, 1977a, p. 10, lo-
cality Aomori- 1). Observed Dec. 1970 by K.
Izawa(1971,p. \95).NE3.
Okuaso vicinity, Arachi-mura, Tsuruga-gun (^
M^gfl^H^^ffe); Fiikiii, HONSHU;
35°35'N, 136°07'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 39: XXV- 17). See Fig. 2A.
Okugataniyama, Kan-noura-cho, Aki-gun ($S
^^M^^T '^\h); Kochi, SHIKOKU;
33°32'N, 134°17'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 22: X-41). See Fig. 2A.
Okugome, Kimitsu-city (^y$TfJ^7K); Chiba,
HONSHU; 35°11'N, 140°02'E; collected 22
Aug. 1998-30 Mar. 1999 by PRIKU staff;
PRIKU, 10 (skeletons only, including 3 with
fragmented skulls). NE120.
Okuike vicinity, Kawachi-mura, Ishikawa-gun
(^JII?P)Rlrt^^)tilftk); Ishikawa, HONSHU;
36°2rN, 136°43'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 40: XXVI-2). See Fig. 2A.
FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES
163
Okumiyama, Ikata-son, Higashiusuki-gun {MS
+^iP#MW^maJ); MiyazakU KYUSHU;
32°32'N, 131°38'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 15: 11-23). See Fig. 2 A.
Okuno. Sasago-mura, Kitatsuru-gun (db|[PlilP
^^^ll#); Yamanashi, HONSHU; 35°35'N,
138°49'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 49: XXXIII-28). See Fig. 2A.
Okuookura vicinity, Kuzuhara-mura, Yama-
gata-gun ( OJ miP«i^^ll A^flfe); Gifiu
HONSHU; 35°35'N, 136°43'E; reported in
questionnaire surxey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 41: XXVIII- 13). See
Fig. 2A.
Okuradake, Yomogita-mura, Higashitsugaru-gun
(^^^IPMffl^A^S); Aomorh HONSHU;
40°58'N, 140°33'E; reported present before
1939 by N. Kuroda (1938, p. 112; 1940. p.
270). Reported present before 1978 by Study
Group on the Present Status of Japanese Mon-
keys (1977a, p. 10).7V£7.
Okusenju. Hirabayashi-mura. Nakakoma-gun {^
&SiP¥tfWa^lljM); YamanashL
HONSHU; 35°34'N, 138°24'E; reported in
questionnaire sur\ey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 48: XXXIII- 10). See
Fig. 2A.
Okutachikawa vicinity. Tanano-son. Katsu-
ura-gun (SMfPai^^aHJH^): Tokushima,
SHikoKU; 33°56'N, 134°31'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 23: XI- 18). See Fig.
2A.
Okutama-machi, Nishitama-gun (?I^01P^^
0 BTT ); Tokyo Metropolis, HONSHU; ca.
35°49'N, 139°06'E; mtDNA samples collected
before 1998 by Y. Kawamoto (1997, p. 33;
2002, p. 60). A^E; 75.
Okutani. Amatsukominato-machi, Awa-gun ($
BIP^^d^^^fflTH^); Chiba, HONSHU:
35°08'N, 140°15'E: collected 3 Aug. 1996 by
PRIKU staff; PRIKU, 1 (skeleton only, with
fragmented skull). NE125.
Okutsugawa National Forest, Hirodo-son, Ka-
tsuta-gun (i^Ba^PiAP^M^JIISW^);
Okayama, HONSHU; 35°09'N, 134°08'E; re-
ported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 27:
XV-22). See Fig. 2A.
Okuuchi-mura. Hata-gun (<#^lPll!*l^); Ko-
chi, SHIKOKU; 33°13'N, 132°53'E; reported
present in 1938 by N. Kuroda (1938, p. 112;
\940, p. 270). SW80.
Okuyama. Kyonan-machi, Awa-gun (^J^IP^
it fflj ^ Lil ); Chiba, HONSHU; 35°07'N,
139°54'E; collected 19 Aug. 1998 by PRIKU
staff; PRIKU. 1 (skeleton only). NE120.
Okuyama, Mitoya-cho. lishi-gun (ISS^PHZIM
HI H Oj ); Shimane. HONSHU; 35°12'N,
132°52'E; collected 21 Nov. 1994 by PRIKU
staff; PRIKU, 1 (skeleton only). SW59.
Okuyama, Nibukawa-mura, Ochi-gun (M^^PM
Jlf^HUl); Ehime. SHIKOKU; 33°58'N,
133°00'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 21:IX-27). SeeFig. 2A.
Okuyamasawa National Forest. Kamikoani-mura,
Kitaakita-gun (db%*(BaiP±'>PqIt^liaj)H
SW^); Akita, HONSHU; 40°00'N, 140°19'E;
reported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 56:
XLIV-6). See Fig. 2A.
Oma-machi. Shimokita-gun ( T db IP A Pal fflJ );
Aomori. HONSHU; ca. 41°32'N. 140°55'E;
mtDNA samples collected before 1998 by Y.
Kawamoto (1997. p. 33). NEl.
Omachi. hills near (Affllrfi) (= Oide, hills near),
HONSHU; ca. 36°30'N, 137°52'E; collected 12
Nov. 1882 by P. L. Jouy; USNM, 1 (specimen
not located, information from USNM field
catalog). 7V£97.
Omachi-city (AffllTtJ); Nagano, HONSHU; ca.
36°30'N, 137°52'E; mtDNA samples collected
before 1999 by Y. Kawamoto (1998, p. 54;
2002, p. 60). NE90.
Ome-city ( ff ^S rfi ); Tokyo Metropolis,
HONSHU; ca. 35°47'N, 139°15'E; mtDNA
samples collected before 1998 by Y. Kawa-
moto (1997. p. 33). 7V£77i.
Omishima. Senzaki-cho, Otsu-gun (A^lP'flljil'^
fflJff^S) (= Oomishima; Oumishima); Ya-
maguchi, OMISHIMA; ca. 34°25'N. 131°11'E;
reported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano. 1974, p. 25:
XIII-5). Occurrence until 1950 reported by K.
Kawanaka (1973, p. 115). population subse-
164
FIELDIANA: ZOOLOGY
quently extinct. SW67.
Onagarayama Forest, Nakano-mura, Minamia-
mabe-gun (]^)§^?P4^if;f^r^^¥aJ^); Oita,
KYUSHU; 32°59'N, 131°48'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 16: III- 12). See Fig.
2A.
Ondayama vicinity, Yunoyama-mura, Onsen-gun
(S^^iP^ULl^i^fflUjffe); Ehime, SHIKOKU;
33°53'N, 132°5rE; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 21: IX-23). See Fig. 2A.
Onigajo, Utsui-son, Toyoura-gun (sM^rt 0
^%Ti0.); Yamaguchi, HONSHU; 34°07'N,
130°58'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 25: XIII- 1). See Fig. 2 A.
Onigajo vicinity, Tateyama-mura, Nakanii-
kawa-gun (4^if JlllPilLil^^^r^ffe); To-
yama, HONSHU; 36°38'N, 137°2rE; reported
in questionnaire survey conducted in 1923 by
K. Hasebe (Iwano, 1974, p. 42: XXIX-5). See
Fig. 2A.
Onoaida, Yaku-cho, Kumage-gun ( M.^W>MP^
^ M ^f^ ); Kagoshima, YAKUSHIMA;
30°15'N, 130°33'E; reported origin of monkeys
kept at Isokoen {W^%'^^''j^M), a park in
Kagoshima City; blood samples were collected
from these captives before 1 992 by Nozawa et
al. (1991, p. 414; 1996, p. 7; T. Shotake,
PRIKU, pers. comm., Nov. 2003). Not
mapped.
Onoaida-Kurio. See Yakushima, Census Area 5.
Onomi and Otaniyama, Matsuyama-mura,
Soo-gun (^J^^^iOl^^rMifm- A^UJ);
Kagoshima, KYUSHU; 31°36'N, 13r06'E;
reported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 12: I-U).
See Fig. 2A.
Onosanrin, Miwa-son, Saeki-gun ('fe'ffi^H^^
Ai?lil#); Hiroshima, HONSHU; coordinates
unknown; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 26: XIV-3). Not mapped.
Onoueyama, Sashiki-mura, Arida-gun (Wffl^S
M >^ ^^ ^ i? ± Lil ); Wakayama, HONSHU;
34°04'N, 135°19'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 33: XX- 1). See Fig. 2A.
Onozumi vicinity, Misaki-mura, Hata-gun (ifil^
IPHllit^^^^ffe); Kochi, SHIKOKU;
32°48'N, 132°52'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 21: X-3). See Fig. 2A.
Oobara, Kisofukushima-machi, Kiso-gun (^e
^^Kt^fifflJAJ^); Nagano, HONSHU;
35°50'N, 137°45'E; collected Nov. 1998 by
PRIKU staff; PRIKU, 1 (skeleton only). NE81.
Ooboke, Sanmyou-son, Miyoshi-gun (H^^H
^ ^ A ^ fe ); Tokushima, SHIKOKU;
33°54'N, 133°45'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 23: XI-20). See Fig. 2 A.
Oobora-Satani, Tara-mura, Yourou-gun (ft^lP
^ ^^ ±MR^"Sy, Gifu, HONSHU;
35°16'N, 136°28'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 40: XXVIIl-4). See Fig. 2 A.
Ooborayama, Ootaki-mura, Chichibu-gun (^^
fP A )t ^^t A )Iql OJ ); Saitama, HONSHU;
35°50'N, 138°53'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 50: XXXVII-2). See Fig. 2A.
Ooda-city (ABSTtT); Shimane, HONSHU; ca.
35°08'N, 132°36'E; mtDNA samples collected
before 2003 by Y. Kawamoto (2002, p. 60).
SW60.
Ooe Takayama, Soshiki-mura, Oochi-gun (b^
lPfi5t^A>IlfUj); Shimane, HONSHU;
35°05'N, 132°29'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 28: XVI-21). See Fig. 2A.
Ooflikasawayama, Shichigashuku-mura, Ka-
tsuta-gun ( XIJ Ba IP ^= ^ =fS ^ A )f )K liJ ); Mi-
yagi, HONSHU; 38°02'N, 140°26'E; reported
in questionnaire survey conducted in 1923 by
K. Hasebe (Iwano, 1974, p. 55: XLIII-3). See
Fig. 2A.
Oogawara, Aiga-mura, Koka-gun (¥M?PI(&/R[
^ A M M ); Shiga, HONSHU; 34°57'N,
136°22'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 38: XXIV-25). See Fig. 2 A.
Ooike beach, Sugari-mura, Kitamuro-gun (it^
mw>mnm^Ammmy, Mie, honshu;
34°06'N, 136°16'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 37: XXIII-43). See Fig. 2A.
FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES
165
Ooiriyama. Sonomura, Kitashitara-gun (db^?^
IPi^AAOj): AichL HONSHU; 35°06'N.
137^44'E; reported in questionnaire sur\ey
conducted in 1923 by K. Hasebe (Iwano. 1974.
p. 40: XXVII-6). See Fig. 2A.
Ooishifukaoawairi \icinit\. Sekitani-mura. Iwa-
t\ine-gun^(^^p^iPil#^AS^JI|Affe):
Migata. HONSHU: 38°04'N. 139°34'E: re-
ported in questionnaire sur\ey conducted in
1923 by K. Hasebe (Iwano. 1974. p. 47:
XXXII-'21). SeeFig. 2A.
Ooiwa. Kimitsu-cit> ( ^ ^ TfT A qs ): Chiba,
HONSHU: 35=12'N. 140°0rE: collected 18
Sep. 1998 and Mar. 1999 by PRIKU staff:
PRIKU. 2 (skeletons only). \E120.
Oojimasan. Takamori-machi. Shimoina-gun (T
^IPiPi^^BrrAlLij): Xagano. HONSHU:
35°34'N. 137^5rE: collected 6 Apr. and 14
May 1998 by PRIKU staff: PRIKU. 2 (skele-
tons only). XE76.
Ooka. Haguro-mura. Niwa-gun (^^?P^M^
'J^E): Aic/iL HONSHU"; ^35°20'N. 136^58'E;
reported absent in questionnaire sur\ey con-
ducted in 1923 by K. Hasebe (Iwano. 1974. p.
40: XXVII-8).
OokancNama. Noma-mura, Yosa-gun (-^^^PS
ral^A^LiJ): Kyoto. HONSHU; 35=40'N.
I35°10'E; reported in questionnaire surxey
conducted in 1923 by K. Hasebe (Iwano. 1974.
p. 35: XXII-5). See Fig. 2 A.
Ookawachiyama \icinit\. Funatsu-mura, Kita-
muro-gun (db^«?P^a'^^^A>R[rt Ojffe); Uie.
HONSHU; 34=13'N. 136°12'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano. 1974. p. 37: XXIII-44). ' See
Fig. 2A.
Ookawamae National Forest, Funaoka-mura.
Kawabe-gun ()qIffllP^p'|5?^AJIIIil SW^);
Akita, HONSHU; 39=38'N. 140=20'E; reponed
in questionnaire sur\ey conducted in 1923 b\
K. Hasebe (Iwano, 1974, p. 56: XLIV-2). See
Fig. 2A.
Ooka\\ara. Ooshika-mura, Shimoina-gun (T#
M^AB^AM^): Xagano. HONSHU;
35=33'N. 138=02'E; collected 12 Apr. 1998 by
PRIKU statT; PRIKU. 1 (skeleton only). XEJO.
Ookawara %icinirv. Neo-mura. Motosu-gun (^
m?P«M^A)qI^ffe); GifiL HONSHU;
35"44'N, 136"34'E; reported in questionnaire
sur\ey conducted in 1923 by K. Hasebe
(Iwano. 1974. p. 41: XXVIII- 10). See Fig. 2 A.
Ookueyama \icinit\. Kitagawa-son, Higa-
shiusuki-gun (^ Q^SPdb'jH*^ AUb UU^);
MiyazakL KYUSHU; 32°44TnJ, I31°30'E; re-
ported in questionnaire sur\ey conducted in
1923 by K. Hasebe (Iwano. 1974. p. 15: 11-27).
See Fig. 2A.
Oomine vicinit}.. Kogurusu. Iruka-mura. Mi-
namimuro-gun (^^-SSPAJ^^d^^^A^
ffe); Mie. HONSHU; 33=52'N. 135°55'E; re-
ported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano. 1974. p. 37:
XXIII-54). See Fig. 2A.
Oomishima. See Omishima.
Oomiya-cho. Watarai-gun (S^IPA^BJ); Mie,
HONSHU; 34°2rN. 136°27E; collected 11
No\. 1997 by PRIKU staff; PRIKU. 1 (skele-
ton only). Slh2.
Oosabiyama vicinitv. Takabayashi-mura,
Nasu-gun (M^mM^^^^MOi^): To-
chigL HONSHU; 37=03'N. 139°50'E; reported
in questionnaire survey conducted in 1923 by
K. Hasebe (Iwano. 1974. p. 51: XXXIX-8).
See Fig. 2A.
Oosaki. Omachi-cit> ( A WJ rfJ A ^ ); Xagano,
HONSHU; 36°29'N. 137°50'E; collected 2
Nov. 1998 by M. Aimi; PRIKU, 2 (skeletons
only). XE90.
Oosato. Onodani-mura, Minamimuro-gun (^^
$?PffiS^^AM); Mie. HONSHU: 33=47?^,
135°59'E; reported in questionnaire survey
conducted in 1923 bv K. Hasebe (Iwano, 1974,
p. 38: XXIII-56). See Fig. 2.\.
Oosawano-machi, Kaminiikawa-gun (_L^JI|?P
A)RSirr); Toyama. HONSHU; ca. 36=34?%',
137°12'E; mtDNA samples collected before
2003 by Y. Kawamoto (2002, p. 60). XE56.
Ooshibayama vicinity. Onikoube-mura, Tama-
zukuri'-gun (3Ei^?P^M^A^aj^); Miyagi,
HONSHU; 38°47'N. 140=40'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano. 1974. p. 55: XLIII-8). See Fig.
2A.
Ooshima-mura. Ooi-gun ( AIS?PA^^); Fukiii,
HONSHU; 35=32^, 135=39'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano. 1974. p. 39: XXV-7). See Fig.
2A.
166
nELDI.\NA: ZOOLOGY
Oosugitani-mura, Taki-gun (^"^^A^^^^);
Mie, HONSHU; 34°16'N, 136°13'E; reported
in questionnaire survey conducted in 1923 by
K. Hasebe (Iwano, 1974, p. 37: XXlII-28). See
Fig. 2A.
Ootani, Makita-mura, Yourou-gun (ft^lP4$![ffl
^A#); Gifu, HONSHU; 35°20'N, 136°28'E;
reported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 40:
XXVIII-6). See Fig. 2A.
Ootaniyama vicinity, Yamanaka-son, Mit-
sugi-gun (^H^OJ^^^A^OJ^); Hi-
roshima, HONSHU; 34°26'N, 133°06'E; re-
ported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 26:
XIV-27). See Fig. 2A.
Ootashiro, Ootaki-machi, Isumi-gun (MP^^A
^* HI AS -ft); Chiba, HONSHU; 35°14'N,
140°10'E; collected 9 Aug. 1997 by PRIKU
staff; PRIKU, 1 (skeleton only). NE126.
Ootsuneki, Tabayama-mura, Kitatsuru-gun (dblrt^
^ m ¥^ 1^\^^ ±n :^); Yamanashi,
HONSHU; 35°48'N, 138°56'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 48: XXXIII-24). See
Fig. 2A.
Oouchi-mura, Igu-gun (PM.W>A\^^ : ^B^
MUL|c^U<^); Miyagi, HONSHU; 37°52'N,
140°50'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 55:XLIII-2). SeeFig. 2A.
Ooyahongou, Ooya-mura, Nima-gun (MB^A
E^AE^^); Shimam, HONSHU; 35°02'N,
132°26'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 28: XVI-27). See Fig. 2A.
Oozakurayama, Kiragawa-mura, Aki-gun ($S
^P^MJII^A^OJ); Kochi, SHIKOKU;
33°20'N, 134°08'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 22: X-45). See Fig. 2A.
Oregon. See Beaverton.
Oshibuchi vicinity, Hobara-mura, Watarai-gun
(Jt^?PaJl^^^)^Jffe); Mie, HONSHU;
34°2rN, 136°4rE; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 37: XXlll-36). See Fig. 2A.
Oshihara-mura, Nima-gun ( S B ?P ^n J^ ^ );
Shimane, HONSHU; coordinates unknown;
reported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 28:
XVI-31). Not mapped.
Oshioi, Takachiho-cho, Nishiusuki-gun (ffiQ^
Wm^%^'Ml^^); Miyazaki, KYUSHU;
32°42'N, 131°18'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 15: 11-31). See Fig. 2 A.
Oshiro, Nukata-cho, Nukata-gun (MBg^P^BaBJ
A -ft ); Aichi, HONSHU; ca. 35°00'N,
137°25'E; blood samples collected before 1992
by Nozawa et al. (1991, p. 414; 1996, p. 6).
NESS.
Osuzu National Forest, Tsuno-cho, Koyu-gun
HPumm^m^m^m^^y, Miyazaki,
KYUSHU; 32°17'N, 131°25'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 15: 11-19). See Fig.
2A.
Ota, Yahata-mura, Kusu-gun (3B^S^1PA<^^A
B9); Oita, KYUSHU; 33°22'N, 13r08'E; re-
ported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 16: III-18).
See Fig. 2A.
Otayama, Iwakura-mura, Arida-gun (WffllP^
^;l^ABgiiJ); Wakayama, HONSHU; 34°03'N,
135°20'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 33:XX-3). SeeFig. 2A. _^
Otoumi, Takahama-cho, Ooi-gun (At5?Pi=i^
^=^M); Fiikiii, HONSHU; ca. 35°32'N,
135°29'E; collected winter 1967 by K. Wata-
nabe (1978, p. 37); PRIKU, 1 (skull only,
mandible missing). SW47.
Otsu-city ( A /* TtT ); Shiga, HONSHU; ca.
35°00'N, 135°52'E; mtDNA samples collected
before 2003 by Y. Kawamoto (2002, p. 60).
SW35.
Otsubo, Kameda, Futtsu-city (S/$rtT%ffl Aif);
Chiba, HONSHU; 35°15'N, 139°52'E; Early
Jomon subfossil (age, ca. 12 Ka) reported be-
fore 2003 by M. Kuroda (2002a, p. 1 15); Wa-
seda University, 1 (mandible only; not seen).
NE119.
Otsukamebora, Nakanoho-mura, Mugi-gun (it
m^^^^^Z.%m); Gifu, HONSHU;
35°34'N, 137°03'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 41: XXVIII-20). See Fig. 2A.
FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES
167
Ouchiyama-mura, Watarai-gun ( i^ ^ ?P A rt ill
^y, Mie, HONSHU; 34°17'N, 136°20'E; re-
ported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 37:
XXIII-29). See Fig. 2A.
Oudake, Kamikuishiki-mura, Nishiyatsu-
shiro-gun (ffiA^t?P± A-fe^ill); Ya-
manashi. HONSHU; 35°31'N, 138°37'E; re-
ported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 49:
XXXIII-33). See Fig. 2A.
Oumishima. See Omishima.
Outaki-mura, Kiso-gun (^H?Pi/^^); Na-
gano, HONSHU^ ca. 35°48'N, 137°33'E;
mtDNA samples collected before 2003 by Y.
Kawamoto (2002, p. 60). NESS.
Ouyama vicinity, Oosugi-mura, Nagaoka-gun
{^^^±Y^¥iM^\hh); KochU SHIKOKU;
33°48'N, 133°43'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 22: X-33). See Fig. 2A.
Owarifuji vicinity, Ikeno-mura, Niwa-gun (^^
iP>fei?WM?is±ffe); Aichi, HONSHU;
35°20'N, 137°0rE; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 40: XXVII- 1 1). See Fig. 2A.
Owase-city ( M M TtJ ); Mie, HONSHU; ca.
34°04'N, 136°12'E; mtDNA samples collected
before 2003 by Y. Kawamoto (2002, p. 60).
SW24.
Oyama vicinity, Shimoukawa-mura, Takeno-gun
(ttg^PT^JII^majfte); Kyoto, HONSHU;
35°44'N, 135°13'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 35: XXII-2). See Fig. 2A.
Oyamada-mura, Ayama-gun (PRllijfPAl-ljffl^);
Mie, HONSHU; caT 34°46'N, 136°14'E;
mtDNA samples collected before 2003 by Y.
Kawamoto (2002, p. 60). SW19.
Oza\\ayama, Sawame-mura, Yamamoto-gun (ill
^ ?P )R g ^ d^ )K LiJ ); Akita, HONSHU;
40°17'N, 140°14'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 56: XLIV-10). See Fig. 2 A.
Ozoechinai. Okuchi-mura, Nomi-gun (BbH^PM
U^MW^^P^); Ishikawa, HONSHU; 36°17'N,
136°38'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 40: XXVI-5). See Fig. 2A.
Rakanyama, Hiromi, Hikimikami-mura,
Mino-gun (ll^^lPE5B±WiAB^:^ > Ul);
Shimam, HONSHU; 34°31'N, 132°00'E; re-
ported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 28:
XVI-12). SeeFig. 2A.
Rebunto (^L:^ft); Hokkaido, REBUNTO; ca.
45°23'N, 141°02E; 2 humeri (artificially modi-
fied) and 1 ulna — all 3 presumably imported to
Rebunto — collected at archaeological site be-
fore 1982 (Nishimoto, 1981, p. 430). Not
mapped.
Rengeiwa vicinity, Mikata-mura, Shisou-gun (5^
M^^y^^W.^^^); Hyogo, HONSHU;
35°irN, 134°36'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 30: XVIII-9). See Fig. 2 A.
Rokujuri, Azuma-mura, Higashitagawa-gun (M
fflJIIiP^^/N + M); Yamagata, HONSHU;
38°3rN, 140°00'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 55: XLII-7). See Fig. 2A.
Rokujuri. Hongou-mura, Higashitagawa-gun {M
BaJH^^M/N+S); Ya/nagata, HONSHU;
38°34'N, 139°48'E; reported in questionnaire
sur\ey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 55: XLII-6). See Fig. 2A.
Rome, Italy; 4r56'N, 12°30'E; population trans-
located from Takasakiyama. Oita Prefecture,
ca. 1977 (Cozzolino et al., 1992, p. 331; Coz-
zolino & Schino, 1998, p. 860). Not mapped.
Rosando, Uchinomi-cho, Shozu-gun ('J^'S.lPrt
M^ ^^^ M ); Kagawa. SHODOSHIMA;
34°3rN, 134°18'E; provisioned group; birth
season, 1958-1962, reported by Kawai et al.
{\967,pp.31,3S).SW68.
Ryogamisan, Ryogami-mura, Chichibu-gun (^
5<riP ft^ tt W 1^ tt UJ ); Saitama, HONSHU;
36°0rN, 138°53'E; reported in questionnaire
sur\ey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 50: XXXVII- 1). See Fig. 2A.
Ryouzen. See Ryozen.
Ryouzenyama. See Ryozenyama.
Ryozen, Ryozen-mura, Date-gun (#]iM?PMlll^
M Oj ); Fiikushima, HONSHU; 37°47'N,
140°38'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 52:XLI-11). SeeFig. 2A.
Ryozen, Seritani-mura, Inukami-gun (:^_blP^
168
FIELDIANA: ZOOLOGY
# ^ M ^llj ); Shiga, HONSHU; 35°14'N,
136°20'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano. 1974,
p. 38: XXIV-20). See Fig. 2A.
Ryozenyama ( M illj lij ); Shiga, HONSHU;
35°17'N, 136°23'E; field study reported by Y.
Sugiyama and H. Ohsawa (1982b, p. 242).
Blood samples collected before 1992 by No-
zawa et al. (1991, p. 414; 1996, p. 6). SW8.
Ryozenyama, Kashiwabara-mura, Sakata-gun
(J^Bg^^aj^^M^lliai); SMga, HONSHU;
35°2rN, 136°25'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 38: XXIV-17). See Fig. 2A.
Ryozenyama, Toriimoto-mura, Sakata-gun (^ffl
^ A M ^ W M ^llj UJ ); Shiga, HONSHU;
35°15'N, 136°19'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 38: XXIV- 19). See Fig. 2A.
Ryujinyama vicinity, Akitsukawa-mura, Nishi-
muro-gun (® #«1P^>»(^$ Jl| ^fltt Uj ftfe);
Wakayama, HONSHU; 33°49'N, 135°27'E;
reported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 33:
XX-14). SeeFig. 2A.
Ryujinyama vicinity, Inari-mura, Nishimuro-gun
(S^ftlP^^WffittUjffe); Wakayama,
HONSHU; 33°45'N, 135°23'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 33: XX- 19). See Fig.
2A.
Ryujinyama vicinity, Kamiakitsu-mura, Nishi-
muro-gun (®^«lP±ti:)tWil4^U4flfe);
Wakayama, HONSHU; 33°45'N, 135°25'E;
reported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 33:
XX-20). See Fig. 2A.
Ryuoka-mura National Forest, Ochi-gun (j^^?P
^^¥iUM^); Ehime, SHIKOKU; 33°57'N,
132°58'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 21:IX-26). SeeFig. 2A.
Ryuozan, Miyauchi, Yawata-son, Mitsugi-gun
(ffllUlPAIf ^^rtfEiOj); Hiroshima,
HONSHU; 34°28'N, 133°05'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 26: XIV-28). See Fig.
2A.
Ryuuzuyama, Tsudani-son, Yamagata-gun (ULj!^
IPIS^^^HMUj); Hiroshima, HONSHU;
34°39'N, 132°26'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 26: XIV-8). See Fig. 2A.
Sabiyama. See Shiobara-mura.
Saburi-mura, Ooi-gun {i^WM^'^%\\^); Fukui,
HONSHU; 35°26'N, 135°33'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 39: XXV-4). See Fig.
2A.
Sagara-mura, Kuma-gun (^^?P^I.^W); Ku-
mamoto, KYUSHU; ca. 32°14'N, 130°48'E;
mtDNA samples collected before 2003 by Y.
Kawamoto (2002, p. 60). SW91.
Sai-mura, Shimokita-gun (TdblP'fc^W); Ao-
mori, HONSHU; 41°26'N, 140°52'E; reported
present before 1939 by N. Kuroda (1938, p.
112; 1940,p. 270).A^£'2.
Saihara-mura, Kitatsuru-gun (dbtPH^PffiJ^^);
Yamanashi, HONSHU; 35°41'N, 139°0rE;
reported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 48:
XXXIII-26). See Fig. 2A.
Sakasegawa vicinity, Itsuki-mura, Kuma-gun (^
BW>^^^J^M}\\i^y, Kimamoto, KYUSHU;
32°24'N, 130°5rE; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 16: IV-7). See Fig. 2 A.
Samuhikiyama, Ooasa-son, Yamagata-gun (|JL|!^
fPAiS^^^^Oj); Hiroshima, HONSHU;
34°47'N, 132°27'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 26: XIV-9). See Fig. 2 A.
Samusawa, Yamanouchi-machi, Shimotakai-gun
(TiS#iPajyrtfflI*)R); Nagano,
HONSHU; 36°43'N, 138°24'E; collected 2
Aug. 1997-21 Aug. 1998 by M. Aimi; PRIKU,
9 (skeletons only, including 5 with fragmented
skulls). NE49.
Samusawayama, Oota-mura, Hienuki-gun (#S
iP;l!^BaW*>RUj); Iwate, HONSHU; 39°23'N,
141°05'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 56: XLV-8). See Fig. 2A.
Sanada-machi, Chiisagata-gun (d^l^iPKEQWI);
Nagano, HONSHU; ca. 36°27'N, 138°20'E;
mtDNA samples collected before 2003 by Y.
Kawamoto (2002, p. 60). NE46.
Sanda, Kimitsu-city ( ^ /$ T|T H BQ ); Chiba,
FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES
169
HONSHU; 35°19'N, 140°04'E; collected Dec.
1998 by PRIKU staff; PRIKU, 1 (skeleton
only). NE128.
Sangayama, Katakaidani-mura, Shimonii-
kawa-gun (TWf JUIPKM^^H® Uj); To-
yama, HONSHU; 36°46'N, 137°31'E; reported
in questionnaire survey conducted in 1923 by
K. Hasebe (Iwano, 1974, p. 42: XXIX-8). See
Fig. 2A.
Sangenyama, Ochiai, Osaka-cho, Masuda-gun
(MBa^Pd^^fflJ^-^HrB^Oj); Gifii, HONSHU;
35°56'N, 137°25'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 41: XXVIII-32). See Fig. 2A.
Sano-goryorin, Sakae-mura, Nishiyatsushiro-gun
{mJ\i^W>^^i^mW^^); Yamanashi,
HONSHU; 35°18'N, 138°29'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 49: XXXIII-37). See
Fig. 2A.
Sanyoushi, Minmaya-mura, Higashitsugaru-gun
(K)igl5HM^^^fflef); Aomori, HONSHU;
41°10'N, 140°26'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 57: XLVI-13). See Fig. 2A.
Sarugababa, Tonoi-son, Toyoura-gun (^y^^Pi^
E^+)-)l:^J\J\); Yamaguchi, HONSHU;
34°16'N, 13r03'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 25: XIII-3). See Fig. 2A.
Sarugamiyama, Manzai-son, Atetsu-gun (Pr[^?P
7T M ^ ?i tt UJ ); Okayama, HONSHU;
34°53'N, 133°32'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 27: XV-5). See Fig. 2 A.
Sarutsubo vicinity, Sada-son, Kimotsuki-gun (if
]i?P^i^^^i¥ftfe); Kagoshima, KYUSHU;
31°06'N, 130°48'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 12: 1-18). See Fig. 2A.
Sasagamine, Kadono-mura, Hikami-gun (7j<±^
Si?W^^ll#); Hyogo, HONSHU; 35°10'N,
135°00'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 30: XVIII-23). See Fig. 2A.
Sasagou vicinity, Mimaki-mura, Kitauwa-gun
(db^^iPISl^^^^ffe); Ehime, SHIKOKU;
33°03'N, 132°40'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 18: IX-5). See Fig. 2A.
Sasayama-cho, Taki-gun ( ^ ^ IP li Ol fflj );
Hyogo, HONSHU; ca. 35°08'N, 135°21'E;
captured alive, arrived at JMC 26 Feb. 1981,
died 16 Apr.-21 May 1981; JMC, 6 (skulls
only). MtDNA samples collected before 2003
by Y. Kawamoto (2002, p. 60). SW39.
Satsuma-cho, Satsuma-gun ( M B ?P ^ JS fflj );
Kagoshima, KYUSHU; ca. 31°55'N, 130°32'E;
mtDNA samples collected before 2003 by Y.
Kawamoto (2002, p. 60). SW93.
Satsuma-gun (MS^); Kagoshima, KYUSHU;
3r54'N, 130°35'E; reported absent in ques-
tionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 12: 1-7). Not mapped.
Seibu-rindou, Kamiyaku-cho, Kumage-gun {W(
^^±Jl^fflTM^^3i); Kagoshima,
YAKUSHIMA; 30°23'N, 130°25'E; collected
Feb.-Mar. 1990 by PRIKU staff; PRIKU, 2
(skeletons only, including 1 with mandible
missing and 1 with fragmented skull). SWIOl.
Sekinosawa, Ikawa-mura, Abe-gun (^io^^JH
^ll;^)R); Shizuoka, HONSHU; 35°12'N,
138°14'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 45: XXX-7). See Fig. 2A.
Senbagatake vicinity, Hiwasa-cho, Kaifu-gun (^
^^BWtefflI=F^<ri^ffe); Tokushima,
SHIKOKU; 33°44'N, 134°29'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 23: XI-7). See Fig.
2A.
Senchougahara, Aoya, Asahi-mura, Masuda-gun
HONSHU; 36°06'N, 137°29'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 41: XXVIII-33). See
Fig. 2A.
Sengatake, Shonai-mura, Suzuka-gun (ipM^Ji
1*1 ^ ill] <r ^ ); Mie, HONSHU; 34°55'N,
136°27'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 36: XXIII-8). See Fig. 2A.
Senzu-goryourin, Kamikawane-mura, Hai-
bara-gun (^J^^±JII^^^M1®13^^); Shi-
zuoka, HONSHU; 35°06'N, 138°08'E; reported
in questionnaire survey conducted in 1923 by
K. Hasebe (Iwano, 1974, p. 42: XXX-3). See
Fig. 2A.
170
FIELDIANA: ZOOLOGY
Seto, Jizoji-mura, Tosa-gun (±'fe?PitkM^^\)'^
F); Kochi, SHIKOKU; 33°43'N, 133°31'E;
reported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 22: X-28).
See Fig. 2A.
Seto, Kinasa-mura, Kamiminochi-gun (JitKi^^
%U^¥iMr)\ Nagano, HONSHU; 36°41'N,
137°59'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 46: XXXI-24). See Fig. 2A.
Seto, Outaki-mura, Kiso-gun (7Ke?Pi/lW/^
F); Nagano, HONSHU; 35°48'N, 137°32'E;
collected 31 Jul.-lO Sep. 1998 by M. Aimi;
PRIKU, 2 (skeletons only). Collected 7-10
Sep. 1998 by KUPRI staff; KUPRI, 2 (skele-
tons only). NESS.
Shakagadake, Shimamoto-mura, Mishima-gun
{^hW>%^¥^W^TU)- Osaka, HONSHU;
34°52'N, 135°38'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 33: XIX-4). See Fig. 2A.
Shakujougadake, Kouchi-mura, Ano-gun {^M.
WM^^^^T^); Mie, HONSHU; 34°49'N,
136°2rE; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 36: XXIII- 1 2). See Fig. 2 A. ^
Shibisan, Takaono-mura, Izumi-gum (tij^K^isi
m if ^ ^ M UJ ); Kagoshima, KYUSHU;
32°02'N, 130°20'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 12: 1-6). See Fig. 2A.
Shichigashuku-machi, Katsuta-gun (XlJB9?P"t'^
?i fflj ); Miyagi, HONSHU; ca. 38°00'N,
140°25'E; mtDNA samples collected before
2003 by Y. Kawamoto (2002, p. 60). NESS.
Shichigodake, Nakama, Yaku-cho, Kumage-gun
{M^^mX^^m^'^^); Kagoshima,
YAKUSHIMA 30°17'N, 130°30'E; collected
27 Jul. 1987 by PRIKU staff; PRIKU, 2
(skeletons only). SfF7 02.
Shichikawa-mura, Higashimuro-gun (^#ft?P
-bJH^); Wakayama, HONSHU; 33°35'N,
135°44'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 34: XX-40). See Fig. 2A.
Shichimensanroku, Motodate-mura, Mi-
namikoma-gun (i^i!S?P^B^-b® lilM);
Yamanashi, HONSHU; 35°24'N, 138°22'E;
reported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 48:
XXXIII-5). See Fig. 2A.
Shichisou-cho, Kamo-gun (iPj^^P-b^fflJ); Gifu,
HONSHU; ca. 35°34'N, 137°06'E; mtDNA
samples collected before 1998 by Y. Kawa-
moto (1997, p. 33; 2002, p. 60). NE60.
Shiga Heights. See Shiga-kogen.
Shiga-A. See Jigokudani.
Shiga-kogen, Shimotakai-gun (Ti^^lP^xMi^
J^) (= Shiga Heights; includes Yokoyugawa
Valley = Shiga-B); Nagano, HONSHU; ca.
36°42'N, 138°29'E; collected 6 Jul. 1962 and
16 Apr. 1963 by PRIKU staff; PRIKU, 2
(skulls only, including 1 with mandible miss-
ing). MtDNA samples collected ca. 1986-1991
by Hayasaka et al. (1991, p. 400). Blood sam-
ples collected before 1992 by Nozawa et al.
(1991, p. 414; 1996, p. 6). External measure-
ments taken before 1997 by Hamada et al.
(1996a, pp. 98, 99). MtDNA samples collected
before 1998 by Y. Kawamoto (1997, p. 33;
2002, p. 60). NE48.
Shigatakiyama vicinity, Tanami-mura, Nishi-
muro-gun (ffi#«?P BQ M*^f .^^Miiaj ^);
Wakayama, HONSHU; 33°30'N, 135°44'E;
reported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 34:
XX-35). See Fig. 2A.
Shigekiyama, Midorisouzu-mura, Mi-
namiuwa-gun (it^^^lP-^iitP^MTKUj);
Ehime, SHIKOKU; 33°00'N, 132°35'E; re-
ported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 18: lX-3).
See Fig. 2A.
Shigira Shinyabakei, Kakiyama-mura, Shi-
moge-gun (T^?P# OJ ^fimj^IP.!)^);
Oita, KYUSHU; 33°23'N, Oril'E; reported
in questionnaire survey conducted in 1923 by
K. Hasebe (Iwano, 1974, p. 16: III- 16). See
Fig. 2A.
Shigumi, Futtsu-city ( g /$ TfT /"^s ^ ); Chiba,
HONSHU; 35°11'N, 139°57'E; collected 19
Jan.-28 Feb. 1998 by PRIKU staff; PRIKU, 3
(skeletons only). NE120.
Shihatsuyama, Yahata-mura, Oita-gun (A'^^
Ri^^mUlU); Oita, KYUSHU; 33°15'N,
131°3rE; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 16:111-13). See Fig. 2A.
FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES
171
Shikakubidani, Takara-son, Katsuura-gun (0M
^^B^^M'M^y, Tokushima, SHIKOKU;
33°59'N, 134°3rE; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 23: XI- 19). See Fig. 2 A.
Shikakumadake, Shimogou-mura, Shimoge-gun
(T^^T®WS^I&); Oita, KYUSHU;
33°25'N, 131°08'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 16: III-14). See Fig. 2A.
Shikakumadake, Yamautsuri-mura, Shimoge-gun
(T^fPUj^^^ESIS); Oita, KYUSHU;
33°25'N, 131°10'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 16: III- 15). See Fig. 2A.
Shikamizawa, Kamiyaku-cho, Kumage-gun (M.
^?P±M^fflTEB)R); Kagoshima,
YAKUSHIMA; ca. 30°20'N, 130°23'E; ob-
served Aug. 1978 by T. Maruhashi (1982, p.
318; cf. Takasaki & Masui, 1984, p. 311).
SWIOI.
Shimakachisanrin vicinity, Katsuragi-mura, Ki-
tamuro-gun (db#«?P^^WftBUj^ftfe);
Mie, HONSHU; 34°07'N, 136°17'E; reported
in questionnaire survey conducted in 1923 by
K. Hasebe (Iwano, 1974, p. 37: XXIII-42). See
Fig. 2A.
Shimashima, Azumi-mura, Minamiazumi-gun
(]^$*?P$»^ft<?); Nagano, HONSHU;
36°irN, 137°47'E; collected 24 Sep.-19 Dec.
1997 by M. Aimi; PRIKU, 5 (skeletons only).
NE94.
Shimatani vicinity, Ryounai-mura, Taki-gun (^
^IPMrt^ft^ftfe); Mie, HONSHU; 34°19'N,
136°15'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 37: XXIII-26). See Fig. 2A.
Shimizu, Ooshika-mura, Shimoina-gun ("F'^ilP
iPAEW)t7K); Nagano, HONSHU; 35°33'N,
138°03'E; collected 26 Mar .-5 Dec. 1998 by
PRIKU staff; PRIKU, 4 (skeletons only).
NE70.
Shimizusawa, Otaki, Toyooka-mura, Kimi-
tsu-gun (^)*?PSI?5^^A)t)f 7K;R); Chiba,
HONSHU; 35°11'N, 139°59'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 50: XXXVI-3). See
Fig. 2A.
Shimizuyama vicinity, Esumi-mura, Nishi-
muro-gun (ffi^ftfP/Hii^)! TROjffe); Wa-
kayama, HONSHU; 33°32'N, 135°37'E; re-
ported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 34:
XX-33). See Fig. 2A.
Shimogou, Hasumi-mura, Oochi-gun (e^^^
^MWT®); Shimane, HONSHU; 34°53'N,
132°37'E; collected 25 Feb. 1995-13 Mar.
1997 by PRIKU staff; PRIKU, 38 (skeletons
only, including 2 with fragmented skulls and 8
with postcranials only). SW62.
Shimohatsugari, Hatsugari-mura, Kitatsuru-gun
( db tP S iP tJ] ^ ^ T ID ^ ); Yamanashi,
HONSHU; 35°36'N, 138°53'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 49: XXXIII-27). See
Fig. 2A.
Shimoichiba, Ooshika-mura, Shimoina-gun (~F
1^3P^AJ^WTT^^); Nagano, HONSHU;
35°34'N, 138°02'E; collected 14 Mar. 1998 by
M. Aimi; PRIKU 1 (skeleton only). NE70.
Shimokawaba vicinity, Yoshino-mura, Na-
gaoka-gun (MI^^P^i^^TJII^ftfe); Kochi,
SHIKOKU; 33°50'N, 133°36'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 22: X-31). See Fig.
2A.
Shimokita, NW. See Oma-machi.
Shimokita, SW. See Wakinosawa vicinity.
Shimokita Peninsula ( ~F db =i^ fe ); Aomori,
HONSHU; 41°09'N, 140°46'E; absence of
congenital limb malformations reported by T.
Shidei et al. (1981, p. 17). MtDNA samples
collected ca. 1986-1991 by Hayasaka et al.
(1991, p. 400). External measurements taken
before 1997 by Hamada et al. (1996a, pp. 98,
99). MtDNA samples collected before 2003 by
Y. Kawamoto (2002, p. 60). NE5.
Shimokitayama-mura, Yoshino-gun (r'^^PT
dbUj^); Nara, HONSHU; 34°0rN, 135°59'E;
reported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 35:
XXI-5). See Fig. 2A.
Shimonaru, Kiyomitsu-mura, Kitauwa-gun (db^
W ?P )t )^ ^ T ^ ); Ehime, SHIKOKU;
33°09'N, 132°36'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 18: IX-6). See Fig. 2A.
Shimonokae vicinity, Izuta-mura, Hata-gun {^
172
FIELDIANA: ZOOLOGY
^ ^ # s ea ^^r T y ip >iite ); kocm,
SHIKOKU; 32°53'N, 132°57'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 21: X-8). See Fig.
2A.
Shimonotani vicinity, Sakauchi-mura, Ibi-gun
(SH^Jt^rt^Ty^flfe); Gifu, HONSHU;
35°34'N, 136°23'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 40: XXVIII-7). See Fig. 2A.
Shimosakuma, Kyonan-machi, Awa-gun {^M
^iSl^fflTTfe^fBl); Chiba, HONSHU;
35°06'N, 139°51'E; collected 17 Jul. 1997-10
Feb. 1999 by PRIKU staff; PRIKU, 11 (skele-
tons only). NE120.
Shimosanji-mura, Hidaka-gun (Si^?PTLi-l£&
^ ); Wakayama, HONSHU; 33°53'N,
135°27'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 33: XX- 11). See Fig. 2A.
Shimoseinaiji, Seinaiji-mura, Shimoina-gun (~F
# 3P ?P )t 1*1 S& 4t T )f rt i?& ); Nagano,
HONSHU; 35°30'N, 137°42'E; collected 7
Oct.-l Nov. 1998 by M. Aimi; PRIKU, 4
(skeletons only). NE73.
Shimoyama (Till); Aomori. See Shimokita Pen-
insula.
Shimoyama, Kushikino-mura, Hioki-gun ( 0 S
15 $ tK if ;t^^t T 111 ); Kagoshima, KYUSHU;
31°43'N, 130°15'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 12: 1-5). See Fig. 2A.
Shimoyama vicinity, Shimomaki-mura,
Mugi-gun (Stm^Ti^^^TUj^); Gifu,
HONSHU; 35°36'N, 136°55'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 41: XXVIII- 17). See
Fig. 2A.
Shindenyama, Abu-son, Kaifu-gun (^pP^PPrIpP
^l/feaUj); Tokushima, SHIKOKU; 33°48'N,
134°39'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 23:XI-5). SeeFig. 2A.
Shinjou vicinity, Mimi-mura, Mikata-gun {=.^5
^:^WifJ±'f-)'JfiUJ#ftfe); Fukui, HONSHU;
35°35'N, 135°56'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 39: XXV- 14). See Fig. 2A.
Shinshiro-city (llf^TfT); Aichi, HONSHU; ca.
34°54'N, 137°30'E; mtDNA samples collected
before 1998 by Y. Kawamoto (1997, p. 33;
2002, p. 60). NE64.
Shinshiu ("(§#1). See Nagano.
Shinyashiki, lijima-machi, Kamiina-gun (_h#BP
?Pt5ftffriffMi(); Nagano, HONSHU;
35°39'N, 137°53'E; collected 12 Nov. 1997 by
M. Aimi; PRIKU, 1 (skeleton only). NE77.
Shinzan, adjacent to Mogami-gun, Nikko-mura,
Akumi-gun {I^M^Umi^^mZ.^^^
^ )^ Ol ); Yamagata, HONSHU; 39°02'N,
140°06'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 55:XLII-9). SeeFig. 2A.
Shinzan, adjacent to Mogami-gun, Oosawa-mura,
Akumi-gun {^m^i^'R^rnhmzm^t
^ )^ UJ ); Yamagata, HONSHU; 38°59'N,
140°0rE; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 55:XLII-8). SeeFig. 2A.
Shinzan, Mizuho-mura, Shinobu-gun (M^^P^X
-f^^Ht)^!!!); Fukushima, HONSHU; 37°45'N,
140°2rE; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 52: XLI-9). See Fig. 2A.
Shiobara-mura, Nasu-gun (JSPM^^^J^W) (=
Sabiyama; Shioyono); Tochigi, HONSHU;
36°58'N, 139°50'E; observed before 1939 by N.
Kuroda (1938, p. 112; 1940, pp. 270, 272).
NEW.
Shiono, Suzaka-city ( ^ ^ Tti im i? ); Nagano,
HONSHU; 36°38'N, 138°21'E; collected 28
Oct. 1998 by M. Aimi; PRIKU, 2 (skeletons
only). NE49.
Shioyono. See Shiobara-mura.
Shippara, Futtsu-city ( S /$ rtT M i^ ); Chiba,
HONSHU; 35°10'N, 139°54'E; collected 23
Dec. 1997-30 Mar. 1999 (excludes two
specimens with unknown collection dates) by
PRIKU staff; PRIKU, 9 (skeletons only).
NEJ20.
Shiragayama, Motoyama-cho, Nagaoka-gun {-B:
^iP^OjfflJSSUj); Kochi, SHIKOKU;
33°48'N, 133°35'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 22: X-32). See Fig. 2A.
Shirahama-cho, Nishimuro-gun (H-^ft^PS*^
HI ); Wakayama, HONSHU; ca. 33°41'N,
135°2rE; onset of provisioning in 1952 re-
FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES
173
ported by Editorial Committee of Nihonzaru
(1977,p. 112).5^m _
Shiraiiama-machi, Awa-gun (^J^^PS^HJ);
Chiba, HONSHU; ca. 34°54'N, 139°54'E; re-
ported present before 1978 by Study Group on
the Present Status of Japanese Monkeys,
1977b, p. 26). MtDNA samples collected be-
fore 1999 by Y. Kawamoto (1998, p. 54).
NE123.
Shiraki, Hatsuki-mura, Isa-gun ('^'te?P^>^^
S ^ ); Kagoshima, KYUSHU; 32°04'N,
130°32'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 12:1-8). See Fig. 2A.
Shirakiyama vicinity, Mizukami-mura,
Kuma-gun (^^?P7K±:|^fi7Klij'te); Kuma-
moto, KYUSHU; 32°22'N, 131°00'E; reported
in questionnaire survey conducted in 1923 by
K. Hasebe (Iwano, 1974, p. 16: IV-6). See Fig.
2A.
Shirakoyama, Kamiyaku-cho, Kumage-gun (^
^lP±M:^Brrajl|aj); Kagoshima,
YAKUSHIMA; 30°27'N, 130°28'E; collected
25 Feb. 1989 by PRIKU staff; PRIKU, 22
(skeletons only, including 1 with fragmented
skull, 1 with mismatched mandible, and 1
postcranials only). SWIOO.
Shiranesan vicinity, Takai-mura, Kamitakai-gun
(±S#lPS#^fi«Ujffe); Nagano,
HONSHU; 36°39'N, 138°26'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 46: XXXI-29). See
Fig. 2A.
Shirataki, Tasuki-son, Toyoura-gun (M/^lPffl^
:^fi^); Yamaguchi, HONSHU; 34°16'N,
131°0rE; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 25:XIII-4). SeeFig. 2A.
Shiratani vicinity, Tokuyama-mura, Ibi-gun (S
HiP^^Uj^e^fte); Gifu, HONSHU;
35°40'N, 136°29'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 40: XXVIII-8). See Fig. 2 A.
Shirayama National Forest, Kamishinjou-mura,
Minamiakita-gun {mXm^±Wi^^ 6 Uj S
W^); Akita, HONSHU; 39°49'N, 140°09'E;
reported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 56:
XLIV-5). See Fig. 2A.
Shiroiwayama, Yamae-mura, Kuma-gun (^S?P
Uj XL ^ S ^ UJ ); Kiimamoto, KYUSHU;
32°18'N, 130°46'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 16: IV-4). See Fig. 2A.
Shirokiyama, Ichikawa-son, Takata-gun (i^BQlP
TfT Jll ;^ fi tK Oj ); Hiroshima, HONSHU;
34°33'N, 132°38'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 26: XIV- 17). See Fig. 2A.
Shirokiyama vicinity, Miiri-son, Asa-gun ($fe
^PHA^S^Ojffe); Hiroshima, HONSHU;
34°34'N, 132°32'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 26: XIV- 14). See Fig. 2A.
Shiroyama, Hatagasako-mura, Kanoashi-gun (M
S.?Ptt0ifi^J^aj); Shimane, HONSHU;
34°28'N, 13r44'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 28: XVI-6). See Fig. 2A.
Shiroyama, Tsuwano-cho, Kanoashi-gun (MS
iP)$^i?fflJ^|ij); Shimane, HONSHU;
34°28'N, 131°45'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 28: XVI-7). See Fig. 2A.
Shizuoka, HONSHU; 34°35'-35°38'N,
137°30'-139°08'E; collected before 2002 by
PRIKU staff; PRIKU, 1 (skeleton only). Not
mapped.
Shobuzaki, Yoshihama-mura, Kesen-gun (^"fllj
iP^^^MSllif ); hvate, HONSHU; 39°09'N,
141°49'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 56: XLV-5). See Fig. 2A.
Shodoshima ( ^J^ S 1 ); Kagawa, SHODO-
SHIMA; 34°23'-34°33'N, 134°07'-134°21'E;
reported present before 1941 by N. Kuroda
(1940, p. 270). MtDNA samples collected
from captive-bom monkeys before 1987 by
Hayasaka et al. (1986, p. 346). Blood samples
collected before 1992 by Nozawa et al. (1991,
p. 414; 1996, p. 6). MtDNA samples collected
before 2003 by Y. Kawamoto (2002, p. 60).
Not mapped.
Shodoshima [1] ( d^ S. ft ); Kagawa,
SHODOSHIMA; ca. 34°31'N, 134°16'E; non-
provisioned, captured 12 Feb. 1962, died in
captivity 6 Apr. 1962; JMC, 1 (skeleton only).
Provisioned, collected 7 Feb. 1981 by Y.
174
FIELDIANA: ZOOLOGY
Hamada; PRIKU, 46 (33 skeletons only, in-
cluding 1 with fragmented skull; 3 skulls with
partial postcranials; 5 skulls only; 5 mandibles
with partial postcranials; and 1 mandible only).
SIV68.
Shodoshima [2] ( ''J'' S A ); Kagawa,
SHODOSHIMA; 34°30'N, 134°18'E; external
measurements taken before 1997 by Hamada
et al. (1996a, pp. 98, 99). SW68.
Shogatake vicinity, Ichinose-mura, Yourou-gun
{^^^-:^M^^tf'^i\k); Gifu, HONSHU;
35°16'N, 136°30'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 40: XXVIII-5). See Fig. 2A.
Shokawagoe vicinity, Kitatonda-mura, Nishi-
muro-gun (M^«?Pdb S ffl ^^tiS JH Mffe);
Wakayama, HONSHU; 33°39'N, 135°26'E;
reported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 34:
XX-25). See Fig. 2A.
Shorenjisan vicinity, Minowa-mura, Naga-gun
(^Mi^Xft^^tffa^lljffe); Me, HONSHU;
34°35'N, 136°07'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 36: XXIlI-16). See Fig. 2A.
Shougenyama, Kamikane-mura, Higashiyama-
nashi-gun (mOj^^Ptt^^^^iilil); Yama-
nashi, HONSHU; 35°47'N, ]38°50'E; reported
in questionnaire survey conducted in 1923 by
K. Hasebe (Iwano, 1974, p. 48: XXXIII-21).
See Fig. 2A.
Shouzawa, Kisofukushima-machi, Kiso-gun (^
MiP7K^^ftfflIIE)R); Nagano, HONSHU;
35°49'N, 137°44'E; collected 10 Dec. 1998 by
PRIKU staff; PRIKU, 1 (skeleton only). NE81.
Shuri Castle, Shuri-Tounokura-cho, Naha-city
(^SPHrtJMM^'MfflTMM^); Okinawa,
OKINAWA; 26°13'N, 127°43'E; mtDNA ex-
tracted in 1999 from skeletal fragments (age ca.
0.4 Ka) by T. Mouri et al. (2000, p. 87); frag-
ments apparently derived from captives intro-
duced from Yakushima. Not mapped.
Sokujitsunomine, Kitakata-mura, Higa-
shiusuki-gun (^Q^l^db^^^^TM 0 ^); Mi-
yazaki, KYUSHU; 32°33'N, 131°27'E; re-
ported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 15: 11-28).
See Fig. 2A.
Soma-city (^l/^rfi); Fukushima, HONSHU; ca.
37°48'N, 140°55'E; mtDNA samples collected
before 2003 by Y. Kawamoto (2002, p. 60).
NE38.
Somakawa-mura, Kamiukena-gun (_L/?!^^tlii
Jll^); Ehime, SHIKOKU; 33°46'N, 133°07'E;
reported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 21:
IX-19). SeeFig.2A.
Sone vicinity, Minamiwauchi-mura, Minami-
muro-gun (l^#g?Pj^li^[^^^^ffe); Mie,
HONSHU; 33°58'N, 136°12'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 37: XXIII-49). See
Fig. 2A.
Soratani, Kasubuchi-mura, Oochi-gun (e^?P^
M^V=P'^); SMmane, HONSHU; 35°05'N,
132°36'E; reported in questionnaire survey
conducted in 1 923 by K. Hasebe (Iwano, 1 974,
p. 28: XVI-22). See Fig. 2A.
Souro vicinity, Shimokawakuchi-mura, Hata-gun
di^l^TJII P^^S-te); Kochi, SHIKOKU;
32°46'N, 132°50'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 21 : X-2). See Fig. 2A.
South Texas. See Arashiyama West.
Sudoyama vicinity, Sudo-mura, Fuji-gun (sdi
15^5$;fvtMytUj'te); Shizuoka, HONSHU;
35°11'N, 138°47'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 45: XXX- 13). See Fig. 2A.
Sugadaira, Sanada-machi, Chiisagata-gun {^b^
^ W g fflj ^ 5|Z ). Magano, HONSHU; ca.
36°31'N, 138°19'E; mtDNA samples collected
before 2003 by Y. Kawamoto (2002, p. 60).
NE47.
Sugawayama vicinity, Nahari-cho, Aki-gun (^
g^^¥^IJfflI^JIiajfte); Kochi, SHIKOKU;
33°24'N, 134°02'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 22: X-47). See Fig. 2A.
Sugino-mura, Ika-gun (##^l!^i?W); Shiga,
HONSHU; 35°33'N, 136°16'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 38: XXIV- 13). See
Fig. 2A.
Sumikawa vicinity, Hikimishimo-mura,
Mino-gun (HJIIPEBT^^JH^); Shimane,
HONSHU; 34°35'N, 131°57'E; reported in
questionnaire survey conducted in 1923 by K.
FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES
175
Hasebe (Iwano, 1974, p. 28: XVI- 1 1). See Fig.
2A.
Sumita-cho, Kesen-gun {^i\hW>i^^^); hvate,
HONSHU; ca. 39°08'N, 141°36'E; mtDNA
samples collected before 1998 by Y. Kawa-
moto (1997, p. 33). A^£7 9.
Sumugitani, Miyajima-mura, Kitakuwada-gun
(db#ffl?P^ft^^^#); Kyoto, HONSHU;
35°15'N, 135°32'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 35: XXII- 15). See Fig. 2A.
Sunada, Amatsukominato-machi, Awa-gun ($
MlP^^d^^JIfflJ^Ba); Chiba, HONSHU;
35°07'N, 140°10'E; collected 7 Mar. 1998-30
Mar. 1999 by PRIKU staff; PRIKU, 19
(skeletons only, including 1 with fragmented
skull). NE125.
Sunomata vicinity, Ukegawa-mura, Higashi-
muro-gun (m^ft^lfJII ^X ^ ^ ^ fife);
Wakayama, HONSHU; 33°48'N, 135°48'E;
reported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 34:
XX-50). See Fig. 2A.
Suzawa, Minamishinano-mura, Shimoina-gun
( T # IP IP m ft )i ^ ^ )H ); Nagano,
HONSHU; 35°17'N, 137°56'E; collected 13
Dec. 1998 by PRIKU staff; PRIKU, 1 (skele-
ton only, with fragmented skull). NE68.
Tabayama-mura, Kitatsuru-gun (db|[PI§?P^/^
\h^)\ YamanashU HONSHU; ca. 35°47'N,
138°56'E; mtDNA samples collected before
2003 by Y. Kawamoto (2002, p. 60). NE114.
Tachibana, Mugi-cho, Kaifti-gun {M^W^^l^
^ ); Tokushima, SHIKOKU; 33°42'N,
134°26'E; collected 18 Mar. 2002 by M. Aimi;
PRIKU, \.SW72.
Tachibanaseiryuu National Forest, Nano-
kawa-mura, Agawa-gun (n"JII?P^i?JIIW^
mUm^^)^ Kochi, SHIKOKU; 33°34'N,
133°08'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 22:X-21). SeeFig. 2A.
Tachigouchi, Muikaichi-mura, Kanoashi-gun (M
£lPASTl?:ftlZ:)RlP«;i); Shimam, HONSHU;
34°2rN, 131°57'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 28: XVI-2). See Fig. 2A.
Tadachi-goryorin, Tadachi-mura, Nishichi-
kuma-gun (S^SiP BQ ll ^BQ ll^3^f^);
Nagano, HONSHU; 35°36'N, 137°33'E; re-
ported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 45:
XXXI-4). See Fig. 2A.
Tadachiokuyama, Ogawa-mura, Kanoashi-gun
( E S: iP ^J^ Jll ^ e ^ H Lil ); Shimane,
HONSHU; 34°30'N, 131°47'E; reported in
questionnaire sur\'ey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 28: XVI-8). See Fig.
2A.
Tadeno, Asakura-mura, Kanoashi-gun (^.S:?P
l^5tW#i?); Shimane, HONSHU; 34°21'N,
131°54'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 28: XVI-3). See Fig. 2A.
Tado Mountain Range, Ishizu-mura, Kaizu-gun
(^^)i?P^)i^^igLilM); Gifu, HONSHU;
35°08'N, 136°39'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 40: XXVIII- 1). See Fig. 2A.
Tado Mountain Range, Shiroyama-mura,
Kaizu-gun {M%^^\h¥i^^\^U); Gifu,
HONSHU; 35°09'N, 136°38'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 40: XXVIII-2). See
Fig. 2A.
Tadosan vicinity, Komi-mura, Kuwana-gun (1^
=glP*ll^^]gajfte); Mie, HONSHU;
35°08'N, 136°36'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 36: XXIII- 1). See Fig. 2 A.
Tadosan vicinity, Tado-mura, Kuwana-gun (H
^IP^JgW^i^lijflfe); Mie, HONSHU;
35°07'N, 136°38'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 36: XXIII-2). See Fig. 2A.
Taga-cho, Inukami-gun (:^_hlP^MfflI); Shiga,
HONSHU; ca. 35°13'N, 136°17'E; mtDNA
samples collected before 2003 by Y. Kawa-
moto (2002, p. 60). SW9.
Tagajo-ato, Tagajo-city (^M^TtT^M^Kl^);
Miyagi, HONSHU; ca. 38°18'N, 141°01'E;
mtDNA extracted from subfossils (age, Nara
period, ca. 1.3 Ka) before 2003 by T.
Agatsuma and M. Ishigami (2002, p. 84);
Okumatsushima Jomon-mura History Museum,
2 (femur, tibia; specimens not seen). NE24.
Tagura, Futtsu-city ( M /¥ rfT BQ ^ ); Chiba,
HONSHU; 35°14'N, 139°57'E; collected 18
176
HELDIANA: ZOOLOGY
Dec. 1998 and 30 Mar. 1999 by PRIKU staff;
PRIKU, 5 (skeletons only). NE120.
Tai-mura, lishi-gun (tS^fPBQ^^); Shimane,
HONSHU; 35°12'N, 132°56'E; reported (as
possibly temporary habitat) in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 28: XVI-35). See Fig. 2A.
Taiheisanrin, Sakai-mura, Shimoniikawa-gun (T
llfJlliPJtW;^¥aJ^); Toyama, HONSHU;
36°57'N, 137°38'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 42: XXIX- 12). See Fig. 2A.
Taikoyama, Tsutsukawa-mura, Yosa-gun (%S}
?PMJII^;*!^i^aj); Kyoto, HONSHU; 35°42'N,
135°15'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 35:XXII-4). SeeFig. 2A.
Taimasan, Ikeda-mura, Shozu-gun (-'J'^S.^Pyfeffl
;^ A Jl^ Uj ); Kagawa, SHODOSHIMA;
34°irN, 134°irE; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 24: XlI-5). See Fig. 2A.
Taipei pet shop, Taiwan; 25°03'N, 121°30'E;
purchased in Oct. 1957 by R. E. Kuntz; USNM,
1 . Not mapped.
Taishakugawa vicinity, Nagato-son, Jinseki-gun
(ttS^*/:iW^lRJI|-w); Hiroshima,
HONSHU; 34°53'N, 133°15'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 27: XIV-36). See Fig.
2A.
Taishakukyo, Tojo-cho, Hiba-gun (i;b^?P^^
fflI^f^llB'5); Hiroshima, HONSHU; 34°52'N,
133°14'E; provisioned group; birth season,
1958-1962, reported by Kawai et al. (1967, pp.
37, 39). SW58.
Takabagamori, Mitsuse-mura, Agawa-gun (ra'JII
^^M^M^'T^); Kochi, SHIKOKU;
33°34'N, 133°22'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 22: X-25). See Fig. 2A.
Takagoyama (SSlJLl) (= Mt. Takago); Chiba,
HONSHU ca. 35°10'-35°15'N, 139°55'-
1 40°00'E; provisioning of group III initiated in
1956-1957 (Suzuki, 1972, pp. 333, 334; Study
Group on the Present Status of Japanese Mon-
keys, 1977b, p. 26). Collected ca. 1961 by S.
Azuma; JMC, 1 (skull only, mandible missing).
Blood samples collected before 1992 by No-
zawa et al. (1991, p. 414; 1996, p. 6). MtDNA
samples collected before 1998 by Y. Kawa-
moto (1997, p. 33). Not mapped.
Takagoyama, Tamaki-mura, Kimitsu-gun (^/$:
IP^^if SUl); Chiba, HONSHU; 35°13'N,
139°57'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 50: XXXVI-2). See Fig. 2A.
Takagoyama A (i^^UjA); Chiba, HONSHU; ca.
35°14'N, 140°01'E; provisioned group; birth
season, 1966, reported by Kawai et al. (1967,
pp. 37, 39). NEI2().
Takagoyama S (i^gOjS); Chiba, HONSHU; ca.
35°12'N, 139°59'E; provisioned group; birth
season, 1959-1966, reported by Kawai et al.
(1967,pp. 37, 38).A^£/20.
Takagoyama Tl (ifgOlTl); Chiba, HONSHU;
35°12'N, 139°59'E; provisioning initiated in
1960 (Suzuki, 1972, p. 333). Subgroup cap-
tured in Mar. 1973, died of disease soon after
capture (Koike & Shimamura, 1988, p. 74);
SU, 96 (84 skeletons only, 12 postcraniais
only). External measurements taken before
1997 by Hamada et al. (1996a, pp. 98, 99).
NE120.
Takahama (S^); Fukui, HONSHU; ca. 35°29'N,
135°33'E; mtDNA samples collected from
captive group tranlocated to PRIKU before
1987 by Hayasaka et al. (1986, p. 346). Blood
samples collected before 1992 by Nozawa et al.
(1991, p. 414; 1996, p. 6). Microsatellite DNA
of laboratory specimen analyzed before 1996
by Domingo-Roura et al. (1997, p. 358). Ex-
ternal measurements taken before 1997 by
Hamada et al. (1996a, pp. 98, 99). MtDNA
samples collected before 2003 by Y. Kawa-
moto (2002, p. 60). SW46. _^
Takahama-cho, Ooi-gun (AtS^Pd^W"); Fukui,
HONSHU; 35°28'N, 135°33'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 39: XXV-3). See Fig.
2A.
Takaharasan vicinity, Tamanyu-mura,
Shioya-gun (Jt#?Pi^^Si^Ujffe); To-
chigi, HONSHU; 36°46'N, 139°51'E; reported
in questionnaire survey conducted in 1923 by
K. Hasebe (Iwano, 1974, p. 51: XXXIX-7).
See Fig. 2A.
Takahata-machi, Higashiokitama-gun (^^II?P
FOODEN AND AIM!: SYSTEMATIC REVIEW OF JAPANESE MACAQUES
177
SfflfflJ); Yamagata, HONSHU; ca. 38°00'N,
140°13'E; mtDNA samples collected before
2003 by Y. Kawamoto (2002, p. 60). NE34.
Takai, Takayama-mura, Kamitakai-gun (JiS^
?PSUj;^S^); Nagano, HONSHU; 36°39'N,
138°2rE; collected 16 Jul. and 27 Oct. 1998
by M. Aimi; PRIKU, 4 (skeletons only). NE49.
Takaike-mura, Higashimuro-gun (^^^-ftlPS/til
^ ); Wakayama, HONSHU; 33°32'N,
135°50'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 34: XX-42). See Fig. 2A.
Takakumadake, Kanoya-cho, Kimotsuki-gun (flf
MiP^MBTTSP?!^); Kagoshima, KYUSHU;
31°33'N, 130°53'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 12: 1-12). See Fig. 2 A.
Takakumadake, Takakuma-mura, Kimotsuki-gun
i^mmm^^m^my, Kagoshima,
KYUSHU; 31°23'N, 130°52'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 12: 1-14). See Fig.
2A.
Takakurayama, Ohsawa-mura, Miyagi-gun ("S"^
^A)R^mt[Uy, Miyagi, HONSHU;
38°18'N, 140°45'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 55: XLIII-7). See Fig. 2A.
Takamurayama, Nyuno-son, Toyota-gun (Wffl
?P A ^ W M UJ ); Hiroshima, HONSHU;
34°28'N, 132°5rE; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 26: XIV-20). See Fig. 2A.
Takane, Asahi-mura, Iwafune-gun (^^p?P^ B
W S ^ ); Niigata, HONSHU; 38°20'N,
139°37'E; collected 15 Oct. 1998 by M. Aimi;
PRIKU, 2 (skeletons only). NE31.
Takanosuyama, Kuba-son, Toyota-gun (sffl^
X^^SM^'M.Shy, Hiroshima, HONSHU;
34°34'N, 132°47'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 26: XIV- 18). See Fig. 2A.
Takanosuyama, Takeni-son, Toyota-gun (SB9?P
1tt^fl|y^Uj); Hiroshima, HONSHU;
34°3rN, 132°45'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 26: XIV- 19). See Fig. 2A.
Takaosan, Hachioji-city (Ai-^Tfii^Mllj); 7b-
kyo Metropolis, HONSHU; 35°37'N, 139°13'E;
provisioned population, translocated from
Shodoshima in Sep. 1957; birth season,
1958-1966, reported by Kawai et al. (1967, pp.
37, 38, 47; cf. Study Group on the Present
Status of Japanese Monkeys, 1977b, p. 23).
NEUO.
Takaosan vicinity, Chikano-mura, Nishi-
muro-gun (ffi#8^]ifii?Wift Mill fife); Wa-
kayama, HONSHU; 33°49'N, 135°37'E; re-
ported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 33:
XX-18). SeeFig. 2A.
Takaosan vicinty, Nagano-mura, Nishimuro-gun
0#«?PMi?WSmajffe); Wakayama,
HONSHU; 33°47'N, 135°28'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 33: XX- 15). See Fig.
2A.
Takara-mura, Minamitsuru-gun (l^fPli^^W);
Yamanashi, HONSHU; 35°34'N, 138°51'E;
reported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 49:
XXXIII-29). See Fig. 2A.
Takarajou vicinity, Awa-mura, Ayama-gun (Pr[
\hmni^¥imB.^^y Mie, HONSHU;
34°45'N, 136°18'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 36: XXIII- 14). S_ee Fig. 2A.
Takasakiyama, Oita-city (A^'^TfTi^lli^lil); Oita,
KYUSHU; 33°15'N, 13r32'E; provisioning
initiated Nov.-Dec. 1952 (Itani et al., 1964, p.
2). Birth season, 1958-1966, reported by
Kawai et al. (1967, pp. 38-39). B virus survey
conducted before 1965 by M. Endo et al. (1964,
p. 173). Captured in Jan. 1968, died 15 Feb.
1968; JMC, 1 (skeleton only). Collected in
1976 by PRIKU staff; PRIKU, 2 (1 skeleton
only, 1 mandible and postcranials only). Killed
by dog in 1984; JMC, 1 (skull only). MtDNA
samples collected ca. 1986-1991 by Hayasaka
et al. (1991, p. 400). Collected before 1991 by
K. Wada; PRIKU, 3 (skeletons only). Col-
lected before 1991 by PRIKU staff; PRIKU, 5
(4 skeletons only, 1 skull only). Blood samples
collected before 1992 by Nozawa et al. (1991,
p. 414; 1996, p. 6). Microsatellite DNA of
laboratory specimen analyzed before 1996 by
Domingo-Roura et al. (1997, p. 358). External
measurements taken before 1997 by Hamada
178
FIELDIANA: ZOOLOGY
et al. (1996a, pp. 98, 99). MtDNA samples col-
lected before 2003 by Y. Kawamoto (2002, p.
60). SIV83.
Takasugi vicinity, Ooyama-mura, Kaminii-
kawa-gun (±lffJlliPAllj^St^^); Toyama,
HONSHU; 36°30'N, 137°22'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 42: XXIX-4). See Fig.
2A.
Takasuyama, Takasu-mura, Sakai-gun (^#?P
li^^^tli^aj); FukuU HONSHU; 36°06'N,
136°05'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 39: XXV- 18). See Fig. 2A.
Takata-mura, Higashimuro-gun (!^#-ft^i^B3
^ ); Wakayama, HONSHU; 33°44'N,
135°54'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 34: XX-47). See Fig. 2A.
Takatoriyama, Minobu-mura, Minamikoma-gun
(lt^S?P#EWJKlXaj); Yamanashi,
HONSHU; 35°22'N, 138°25'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 48: XXXIII-4). See
Fig. 2A .
Takatoriyama National Forest, Takaichi-mura,
Takaichi-gun (i^Tt]fPi^T^:ttS15ililSW^);
Nara, HONSHU; 34°28'N, 135°50'E; reported
in questionnaire survey conducted in 1923 by
K. Hasebe (Iwano, 1974, p. 35: XXI-13). See
Fig. 2A.
Takatoriyama National Forest, Takatori-cho,
Takaichi-gun (if TfT^Pif IJ^ffllSUUjaW^);
Nara, HONSHU; 34°25'N, 135°48'E; reported
in questionnaire survey conducted in 1923 by
K. Hasebe (Iwano, 1974, p. 35: XXI-12). See
Fig. 2A.
Takatsuki-cho, Ika-gun (##?PIf >^ BJ); Shiga,
HONSHU; ca. 35°28'N, 136°15'E; mtDNA
samples collected before 2003 by Y. Kawa-
moto (2002, p. 60). SW6.
Takayama, Kuchi-son, Asa-gun (^jilP^Xitfe^
If llj ); Hiroshima, HONSHU; 34°3rN,
132°24'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 26:X1V-15). SeeFig. 2A.
Takeno-cho, Kinosaki-gun ( ^ lli$ IP t^ i? ffll );
Hyogo HONSHU; ca. 35°35'N, 134°44'E; pro-
visioned group, translocated from Kochi be-
fore 1966; birth season, 1965-1966, reported
by Kawai et al. (1967, pp. 38, 47). Collected
ca. 27 Oct. 1971 by JMC staff; JMC, 1 (skele-
ton only). SW49.
Takidani vicinity, Ootaki-mura, Inukami-gun (^
±fP;'^)i^il#fe); Shiga, HONSHU;
35°18'N, 135°56'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 38: XXIV-22). See Fig. 2A.
Takigoshi, Outaki-mura, Kiso-gun (^KelPi/^
^ )t ® ); Nagano, HONSHU; 35°49'N,
137°27'E; collected 4 and 6 Sep. 1998 by M.
Aimi; PRIKU, 2 (skeletons only). NE84.
Takinozawa, Seinaiji-mura, Shimoina-gun (T"^
M^^\^^¥SM(D)R); Nagano, HONSHU;
35°30'N, 137°40'E; collected 5 Dec. 1998 by
PRIKU staff; PRIKU, 1 (skeleton only). NE73.
Takitani, Sugawa-mura, Kanoashi-gun (^JE?P
^JIW)t#); Shimane, HONSHU; 34°33'N,
131°52'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 28:XVI-10). SeeFig. 2A.
Takitani vicinity, Kuda-mura, Atago-gun (^S
W>X^¥iM"^^); Kyoto, HONSHU; 35°15'N,
135°49'E; reported in questionnaire survey
conducted in 1 923 by K. Hasebe (Iwano, 1 974,
p. 36:XXII-21). SeeFig. 2A.
Takiyama, Kitayoshino-son, Katsuta-gun (Mffl
iP db ^ i? ^ >t Ol ); Okayama, HONSHU;
35°06'N, 134°09'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 27: XV-21). See Fig. 2A.
Takiyamagawa vicinity, Kake-cho, Yama-
gata-gun {\hWkWM\^M\^]\\y^^y ^^
ffe); Hiroshima, HONSHU; 34°37'N, 132°21'E;
reported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 26:
XIV-7). See Fig. 2A.
Takuwa, Mitoya-cho, lishi-gun (ISS5PHZJM
fflJ^^^); Shimane, HONSHU; 35°15'N,
132°52'E; collected 12 Dec. 1994-27 Jul. 1997
by PRIKU staff; PRIKU, 8 (skeletons only, in-
cluding 1 with fragmented skull and 1 with
fragmented mandible). SW59.
Tanegashima ( S •? S ); Kagoshima,
TANEGASHIMA; ca. 30°36'N, 130°59'E; oc-
currence in 1905 reported by local residents to
M. P. Anderson (in Thomas, 1906 ["1905"], p.
362). Occurrence in 1940s reported by K. Ka-
FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES
179
wanaka (1973, p. 1 15). Extinction or near ex-
tinction of island population reported by local
residents in April 1970 (Azuma, 1972, p. 262).
SW99.
Taneeawayama vicinity, Funaki-mura, Nii-gun
( ^ M ?P ^p^ ^ ^ S ^ Jll UJ fte ); Ehime,
SHIKOKU; 33°55'N, 133°20'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 21: IX- 16). See Fig.
2A.
Taninoshiriyama vicinity, Kamiiritsu-mura, Mi-
namiamabe-gun CMM^^JiX^M^^^^
Ojfe); Oira, KYUSHU; 32°52'N. 131°56'E;
reported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 16: I1I-9).
See Fig. 2A.
Tano, Handa-mura, Kusu-gun (3E^S^lPtSffl^EB
W)\ Oita, KYUSHU; 33°09'N, 131°15'E; re-
ported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 16: III- 19).
See Fig. 2A.
Tanosawa vicinity, Uchigata-mura, Ki-
tatsugaru-gun (db^^^lP 1*1 ^^ BB ^ >R fife);
Aoniori, HONSHU; 41°03'N, 140°29'E; re-
ported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 57:
XLVI-10). SeeFig. 2A.
Tanzawayama, Kiyokawa-mura, Aikou-gun (S
¥?P>f JII^:l^>RLil); Kanagawa, HcJnSHU;
ca. 35°28'N, 139°10'E; mtDNA samples col-
lected before 1999 by Y. Kawamoto (1998, p.
54). NE109.
Tara-dake, Fujitsu-gun (R^?P^MS); Saga,
KYUSHU; ca. 32°59'N, 130°06'E; observed in
1940s by local residents (Sakura, 1976, p. 151);
subsequently extinct at this locality (Ikeda and
Eguchi, 1978, p. 59). 5PF57.
Tarekaya National Forest, Yuki-cho, Jinseki-gun
{^^W/^^^^ U:^i7SW^); Hiroshima,
HONSHU; 34°47'N, 133°18'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 26: XIV-33). See Fig.
2A.
Tarogawa, Kawasaki-machi, Shibata-gun (^ffl
IP Jll *$ fflj ;!!; HP Jll ); Miyagi, HONSHU;
38°14'N, 140°35'E; observed Dec. 2002-Jan.
2003 by K. Izawa, T. Uno, and H. Fujita (2003,
p. \^).NE25.
Tarudaira vicinity, Horado-mura, Mugi-gun (^
'B.^Mf^^^ )l^^)\ Gifii, HONSHU;
35°37'N, 136°5rE; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 41: XXVIII- 15). See Fig. 2A.
Tanisawayama vicinity, Hikawa-mura, Nishi-
tama-gun (^^MW^]\\^M)R\U^)\ Tokyo
Metropolis, HONSHU; 35°47'N, 139°05'E;
reported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 49:
XXXI V-3). See Fig. 2 A.
Tashida vicinity, Hatta-mura, Inabe-gun (M#?P
>^Ba^^^^Baftfe); Mie, HONSHU; 35°08'N,
136°28'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 36: XXIII-3). See Fig. 2A.
Tashiro, Ootaki-machi, Isumi-gun (MP^?PA^
S BTT BQ ft ); Chiba, HONSHU; 35°13'N,
140°12'E; collected 16 Aug. 1997 by PRIKU
staff; PRIKU, 1 (skeleton only). NE126.
Tashiyama, Murou-mura, Uda-gun (^PS?P^^
^ A 6f Ol ); Nara, HONSHU; 34°31'N,
136°03'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 35:XXI-10). SeeFig. 2A.
Tasmania. See Launceston.
Tatai, Horigane-mura, Minamiazumi-gun (]^$
#iP^^WBa^#); Nagano, HONSHU;
36°18'N, 137°50'E; collected 9 Nov. 1997 by
M. Aimi; PRIKU, 1 (skeleton only). Collected
20 Nov. 1997 and 27 Nov. 1999 by PRIKU
staff; PRIKU. 2 (skeletons only). NE93.
Tataki vicinity, Tokuda-mura. Shusou-gun (j^lil
IP ^> Ba ^ ea >i ffe ); EJume, SHIKOKU;
33°54'N. 133°00'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 18: IX-8). See Fig. 2A.
Tatsufusayama vicinity, Higashimera-son,
Koyu-gun (iPo^^^m^KM^HMLUffe); M-
ya-aki, KYUSHU; 32°15'N, 13ri7'E; re-
ported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 15: 11-16).
See Fig. 2A.
Tatsuka\\ ayama, Sumino-mura, Nii-gun (^M?P
^^^^aUIIOj); Ehime, SHIKOKU; 33=52?^,
133°23'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 21: IX- 14). SeeFig. 2 A.
Tatsuyama-mura, Iwatagun (#ffl?PilU4^);
Shizuoka. HONSHU; ca. 34°52'N, 137°49'E;
180
HELDIANA: ZOOLOGY
mtDNA samples collected before 1998 by Y.
Kawamoto (1997, p. 33; 2002, p. 60). NE66.
Tazawa, Takamori-machi, Shimoina-gun (T"^
IP ?P if ^ BTT Ba )R ); Nagano, HONSHU;
35°35'N, 137°51'E; collected 11 Dec. 1997 by
PRIKU staff; PRIKU, 3 (skeletons only, in-
cluding 1 with fragmented skull). Collected 3
Nov. 1998 by M. Aimi; PRIKU, 4 (skeletons
only). NE76.
Tedayama vicinity, Togo-son Higashiusuki-gun
(maff^m^^^BgOlffe); Miyazakt,
KYUSHU; 32°22'N, 131°28'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 15: 11-20). See Fig.
2A.
Tenjinbara, Takayama-mura, Kamitakai-gun (_h
S#^ifaj;Ht^ttJI); Nagano, HONSHU;
36°41'N, 138°24'E; collected 11 Aug.-7 Sep.
1998 by M. Aimi; PRIKU, 5 (skeletons only).
NE49.
Tenjinkyo, Yoshii-cho, Shituki-gun {^HW^^
ffll^#ll!^); Okayama, HONSHU; 34°39'N,
133°25'E; mtDNA samples collected
1998-2000 by I. Yoshimi and H. Takasaki
(2003, p. 71). 5^56.
Tenjinyama, Yuno-son, Kawakami-gun (Jl|_h^
^ IF W ^ ^ UJ ); Okayama, HONSHU;
34°52'N, 133°25'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 27: XV-4). See Fig. 2A.
Tennoji ( ^ 3E ^ ); Osaka, HONSHU; ca.
34°40'N, 135°30'E; living captive purchased at
market 10 Jun. 1826 by P. F. von Siebold
(Holthuis & Sakai, 1970, p. 67); specimen
possibly included among those sent by von
Siebold to RMNH. Not mapped.
Tenryu-city (^"^Tf]); Shizuoka, HONSHU; ca.
34°53'N, 137°49'E; blood samples collected
before 1992 by Nozawa et al. (1991, p. 414;
1996, p. 6).yV£'66.
Texas. See Arashiyama West.
Tochiyama vicinity, Akaba-mura, Kitamuro-gun
(db#«lP#^^^Ujfte); Mie, HONSHU;
34°14'N, 136°16'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 37: XXIII-41). See Fig. 2A.
Tochu vicinity, Hongawa-mura, Tosa-gun (zhfe
?P ^ Jll ^ F 4^ -te ); Kochi, SHIKOKU;
33°44'N, 133°20'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe (Mito,
1989, p. 25). See Fig. 2A.
Todai. See Nagata-todai.
Toei-cho, Kitashitara-gun (dblS^^P^^BO");
Aichi, HONSHU; ca. 35°02'N, 137°45'E;
mtDNA samples collected before 1998 by Y.
Kawamoto (1997, p. 33; 2002, p. 60). NE67.
Togakushiyama vicinity, Togakushi-mura,
Kamiminochi-gun (±7K[*l?PFH:|4p|iiiJL|
fife); Nagano, HONSHU; 36°43'N, 138°05'E;
reported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 46:
XXXI-25). See Fig. 2A.
Toi-misaki, Kushima-city ( $ Pb^ TfJ |[P ^ ill? ) (=
Cape Toi, Kushima); Miyazaki, KYUSHU;
31°22'N, 131°20'E; provisioned group; birth
season,1966, reported by Kawai et al. (1967.
pp. 37, 39). Collected 31 Dec. 1966 by JMC
staff; JMC, 1 (skeleton only). SW98.
Tojiroyama, Tomisato-mura, Nishiyatsu-
shiro-gun OA-ftlPsM^^'ftUj); Yama-
nashi, HONSHU; 35°26'N, 138°29'E; reported
in questionnaire survey conducted in 1923 by
K. Hasebe (Iwano, 1974, p. 49: XXXIII-35).
See Fig. 2A.
Tokio. See Tokyo.
Tokio Market. See Tokyo market.
Tokiwa, Oomachi-city (AffllTfi^^); Nagano,
HONSHU; 36°29'N, 137°50'E; collected 8 Apr.
1998 by M. Aimi; PRIKU, 1 (skeleton only).
Collected 2 Nov. 1998 by PRIKU staff;
PRIKU, 2 (skeletons only). NEW.
Tokiyama, Toki-mura, Yourou-gun (ft^iPB^^
B^lll); Gifu, HONSHU; 35°14'N, 136°26'E;
reported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 40:
XXVIII-3). See Fig. 2A.
Tokuji, Ichihara-city ( TtT i^ TfT ^> E ); Chiba,
HONSHU; 35°18'N, 140°09'E; collected 5 Feb.
and 30 Mar. 1999 by PRIKU staff; PRIKU, 2
(skeletons only). NE127.
Tokusa, Hagiwara, Oofuji-mura, Higashiyama-
nashi-gun (mOJ^^P AM^i^J^^M); Ya-
manashi, HONSHU; 35°40'N, 138°49'E; re-
ported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 48:
XXXIII-22). See Fig. 2A.
Tokuzen-nishi vicinity, Nishiiyayama-son,
Mima-gun (M^l^Pem^Oj^^^Sffiftfe);
FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES
181
Tokushima, SHIKOKU; 33°54'N, 133°50'E;
reported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 23:
XI-21). SeeFig. 2A.
Tokyo (^^tP) (= Jedo; Tokio; Yedo); Tokyo
Metropolis, HONSHU; 35°42'N. 139°46'E;
obtained 1874-1876 by Mr. v. Hilgendorf (cf.
Schweyer, 1909, p. 8), ZMB, 6 (1 skeleton
only, 5 skulls only). Purchased 1900-1901 by
K. A. Haberer (cf. Schweyer, 1909, p. 8); mu-
seum unknown, 2 (skulls only, possibly in-
cluded among unlocalized specimens exam-
ined). Not mapped.
Tokyo market. (M^f^^) (= Tokio Market);
Tokyo Metropolis, HONSHU; 35°42'N,
139°46'E; purchased 31 Dec. 1882 and 1 1 Feb.
1883 by P. L. Jouy; USNM, 3 or 4 (2 male
skulls, 1 female skull, and 1 female skin [mis-
matched with male skull]). Not mapped.
Tomikawayama, Ooishi-mura, Kurita-gun (^^
?PA^^sJIIUJ); Shiga, HONSHU; 34°52'N,
135°53'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 39: XXIV-28). See Fig. 2A.
Tomogashima, Wakayama-city ( ^ 1^ |1| rfi" S 'It
My, Wakayama, TOMOGASHIMA; 34°17'N,
135°00'E; monkey park population translo-
cated from Yakushima, opened in 1963, closed
in 1972 (cf Shidei et al., 1981, p. 17). SWS2.
Toragamine Mountain Range vicinity, Kurusu-
kawa-mura, Nishimuro-gun (ffi#S^^ffiJI|
^M.T^\\\U^); Wakayama, HONSHU;
33°5rN, 135°28'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 33: XX-16). See Fig. 2A.
Torihama shell mound, Mikata-cho, Mikata-gun
{^15^^15^ M'ikRW); Fukui, HONSHU;
coordinates unknown; mid-Jomon subfossils
(age, 5-6 Ka) reported by M. Kuroda (2002a,
p. 118); Jakuo Historical and Ethnographic
Museum, 2 (skulls only; not seen). Not
mapped.
Toriidoyama, Chikusa-mura, Mie-gun (HMIP^
SWBMpUj); Mie, HONSHU; 35°02'N,
136°29'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 36: XXIII-4). See Fig. 2A.
Torishima, Kushima-city (^PalrfiBM); Miya-
zah, TORISHIMA; 3r27'N, 131°24'E; a
young male monkey swam ca. 600 m to this
island in 1970 (Mito, 1980, p. 29). SW97.
Totorogauchi, Kitagou-son, Higashiusuki-gun
(mQttlPdb0W±<? S<rrt); Miyazaki,
KYUSHU; 32°31'N, 131°27'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 15: 11-24). See Fig.
2A.
Totorogauchi vicinity, Kadokawa-son, Higa-
shiusuki-gun (mQ^?PP^JI|^±<? S^rt
ffe); Miyazaki, KYUSHU; 32°28'N, 131°38'E;
reported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 15: 11-22).
See Fig. 2A.
Totsukawa-mura, Yoshino-gun ( n^if ?P~I~/$JI|
^); Nara, HONSHU; 34°00'N, 135°45'E; re-
ported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 35:
XXl-4). See Fig. 2A.
Towada, Houokusawa-mura, Kamikita-gun (Ji
db?P^^||)R + ^ffl); Aomori, HONSHU;
40°33'N, 141°03'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 57: XLVI-18). See Fig. 2A.
Towadako Lake vicinity, Nanataki-mura, Ka-
zuno-gun (E^ ^-b )i W + ^ BB /^-f^M);
Akita, HONSHU; 40°05'N, 140°34'E; reported
in questionnaire survey conducted in 1923 by
K. Hasebe (Iwano, 1974, p. 56: XLIV-[13]).
See Fig. 2A.
Toyamagoryouchi vicinity, Nagashino-mura,
Minamishitara-gun (Mixi^^:1^^5gUj1fil
^^^);Aichi, HONSHU; 34°55'N, 137°37'E;
reported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 40:
XXVII-2). See Fig. 2A.
Tsubaki Wild Monkey Park, Shirahama-cho,
Nishimuro-gun O^ftlPS^fflJ^^m^A^
% ); Wakayama, HONSHU; 33°36'N,
135°24'E; positional behavior studied
1995-1997 by K. Chatani (2003, p. 14).
MtDNA samples collected before 2003 by Y.
Kawamoto (2002, p. 60). SW29.
Tsubakiyama vicinity, Ookouchi-mura, Nishi-
yatsushiro-gun (liA^ftlP AMrt^lSUl'te);
Yamanashi, HONSHU; 35°24'N, 138°30'E;
reported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 49:
XXXIII-36). See Fig. 2A.
182
FIELDIANA: ZOOLOGY
Tsubayama, Ikegawa-cho, Agawa-gun (n-JUlP
)teJI|fflI^Lij); Kochi, SHIKOKU; 33°39'N,
133°irE; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 22: X-22). See Fig. 2A.
Tsuchikurayama, Kamiarusu-mura, Kesen-gun
(^flljlP±W'(±;^±5taj); Iwate, HONSHU;
39°28'N, 141°26'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 56: XLV-3). See Fig. 2A.
Tsuchiosanrin vicinity, Ishigasato-son,
Atetsu-gun (PqI^?P^g^^±MLJj#fte);
Okayama, HONSHU; 34°54'N, 133°30'E; re-
ported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 27: XV-6).
See Fig. 2A.
Tsuchiyama-cho, Koka-gun [1] (¥M?P±ajfflI);
Shiga, HONSHU; ca. 34°57'N, 136°17'E;
mtDNA samples collected before 2003 by Y.
Kawamoto (2002, p. 60). SW15.
Tsuchiyama-cho, Koka-gun [2] (¥M?P±UjfflI);
Shiga, HONSHU; ca. 34°57'N, 136°19'E; col-
lected Dec. 1980 by TPM staff; TPM, 1 (skull
only). SW15.
Tsugaru. See Nishimeya-mura, Minamit-
sugaru-gun.
Tsugaru-touge, Ajigasawa-machi, Nishitsu-
garu-gun (ffi)$^?Pii <r >RfflJ)$g|lt#); Ao-
mori, HONSHU; 40°34'N, 140°09'E; mtDNA
samples collected before 2003 by Y. Kawa-
moto (2002, p. 60). NE9.
Tsukagoshi, Ootaki-machi, Isumi-gun (MP^lPA
^M^MM); Chiba, HONSHU; 35°14'N,
140°09'E; collected 31 Jul. 1997-23 Nov. 1997
by PRIKU staff; PRIKU, 5 (skeletons only, in-
cluding 1 with fragmented skull). NE126.
Tsukinoya, Kisuki-cho, Ohara-gun (AJ^IP^/^
fflJ^yM); Shimane, HONSHU; 35°13'N,
132°56'E; collected 21 Nov. 1994 by PRIKU
staff; PRIKU, 3 (skeletons only, including 1
with fragmented skull and 1 with mandible and
postcranials only). SW59.
Tsukiyama vicinity, Fujiwara-machi, Shioya-gun
(^^^SJ^fflTv^Ujffe); Tochigi, HONSHU;
36°50'N, 139°39'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 51: XXXlX-5). See Fig. 2A.
Tsukiyodaira, Takamori-machi, Shimoina-gun
{T ^M^nm^ n^^y, Nagano,
HONSHU; 35°34'N, 137°52'E; collected 13
Mar.^ Aug. 1998 by M. Aimi; PRIKU, 14
(skeletons only). NE76.
Tsukiyonodan, Mutsuai-mura, Minamikoma-gun
(lt&0?PBi-^^.^^IS:); Yamanashi,
HONSHU; 35°14'N, 138°23'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 48: XXXIII-2). See
Fig. 2A.
Tsukizaki, Ichihara-city ( rfT J^Tfi ^ lli^); Chiba,
HONSHU; 35°18'N, 140°08'E; collected 21
Jan.-30 Mar. 1998 by PRIKU staff; PRIKU, 6
(skeletons only). NE127.
Tsukubasan, Tsukuba-machi, Tsukuba-gun (m^^
IP ^ }:^ fflj ^ /:^ U4 ); Ibaraki, HONSHU;
36°14'N, 140°08'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 51: XXXVIII-1). See Fig. 2A.
Tsukui-machi, Tsukui-gun (y^^^^/$^#ffll);
Kanagawa, HONSHU; ca. 35°31'N, 139°16'E;
mtDNA samples collected before 1998 by Y.
Kawamoto (1997, p. 33; 2002, p. 60). NE108.
Tsunagi, Yamagata-mura, Kunohe-gun {%WW^
UU ff^ ^ 1^ ); Iwate, HONSHU; 40°10'N,
141°39'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 56:XLV-11). SeeFig. 2A.
Tsuneroku vicinity, Tomiyama-mura, Hata-gun
(<#^iPaUj^^7Nffe); Kochi, SHIKOKU;
33°07'N, 133°00'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 21: X-14). See Fig. 2 A.
Tsunogawa-Akana vicinity, Mitani-mura, Taka-
shima-gun (^ft?PH^;f^r^JI^^^=Sflfe); 5%fl,
HONSHU; 35°25'N, 135°55'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 38: XXIV-7). See Fig.
2A.
Tsunotani, Tsuga-mura, Oochi-gun (e^^PfPM
;1^ ^ ^ ); Shimane, HONSHU; 34°58'N,
132°38'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 28: XVI-25). See Fig. 2A.
Tsuriganeyama, Nanaori-son, Nishiusuki-gun (®
e^^-biff^t^Mlij); Miyazaki, KYUSHU;
32°4rN, 131°27'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 15: 11-29). See Fig. 2A.
Tsuro vicinity, Kaminada-mura, Hata-gun (<#^
FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES
183
?P ± ^ W )i S fte ); Kochi, SHIKOKU;
32°45'N, 133°0rE; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 21: X-7). See Fig. 2A.
Tsutsumori, Ootaki-machi, Isumi-gun (MP^^A
^gffllM0); Chiba, HONSHU; 35°13'N,
140°09'E; collected 30 Jun. 1997 and 6 Aug.
1998 by PRIKU staff; PRIKU, 2 (skeletons
only). NE126.
Tsutsumori National Forest, Oikawa-mura,
Isumi-gun (HP^i^^JII^M^aW^);
Chiba, HONSHU; 35°11'N, 140°12'E; re-
ported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 50:
XXXVI-7). See Fig. 2A.
Tuino. See Jinrio.
Uchiyama, Yoshinaga-mura, Fuji-gun (gdr^'o
^Wl^lLiJ); Shizuoka, HONSHU; 35°14'N,
138°46'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 45: XXX- 12). See Fig. 2 A.
Ueda, Hasumi-mura, Oochi-gun (b^^^MH
W ± Ba ); Shimane, HONSHU; 34°51'N,
132°41'E; collected 2 Jul. 1996-12 Nov. 1997
by PRIKU staff; PRIKU, 29 (skeletons only).
SW62.
Ui, Ishigaki-mura, Arida-gun (WBB^^ilW^
# ); Wakayama, HONSHU; 34°00'N,
135°20'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 33:XX-2). SeeFig. 2A.
Ujitahara-mura, Tsuzuki-gun (^§^?P^/p BBJ^
^); Kyoto, HONSHU; 34°50'N, 135°52'E;
reported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 36:
XXII-29). See Fig. 2A.
Umegatani vicinity, Miuchi-mura, Onsen-gun
{iB^^^^P^^^T'^^); Ehime, SHIKOKU;
33°47'N, 132°57'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 21: IX-25). See Fig. 2A.
Unazuki-machi, Shimoniikawa-gun (Tl!f JII?P^
^^^fflj); Toyama, HONSHU; ca. 36°51'N,
137°34'E; mtDNA samples collected before
2003 by Y. Kawamoto (2002, p. 60). NE53.
Unsen. See Kyogatake.
Urayama, Shino-mura, Minamikuwada-gun (]^
^Ba^P^^ROl); Kyoto, HONSHU; 34°59'N,
135°38'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 36: XXlI-25). See Fig. 2A.
Ushigadake, Ikazawa-mura, Minamiuonuma-gun
(|lg^)S?PE + )R*t4^^); Niigata,
HONSHU; 37°03'N, 138°57'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 47: XXXlI-5). See
Fig. 2A.
Ushimaki, Takamori-machi, Shimoina-gun (T#
M^M^^ ^^); Nagano, HONSHU;
35°34'N, 137°51'E; collected 11 Jan. 1998 by
M. Aimi; PRIKU, 1 (skeleton only). Collected
9 Mar.-8 Dec. 1998 by PRIKU staff; PRIKU,
27 (skeletons only, including 3 with frag-
mented skulls). NE76.
Ushiokuyama vicinity, Okunoda-mura, Higashi-
yamanashi-gun (^Ul^^PEif BgW^HOj-
w ); Yamanashi, HONSHU; 35°41'N,
138°49'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 48: XXXlII-23). See Fig. 2A.
Ushiroyama, Hasumi-mura, Oochigun (b^?P
^^^Wik\h); Shimane, HONSHU; 34°52'N,
132°38'E; collected 19 Dec. 1995-29 Jul. 1998
by PRIKU staff; PRIKU, 15 (skeletons only).
Collected 11 Mar. 1999-14 Jul. 2000 by M.
Aimi; PRIKU, 21 (skeletons only). SW62.
Ushitakisan, Yamataki-mura, Senboku-gun {^
db^Uj>i^^)1iaj); Osaka, HONSHU;
34°24'N, 135°28'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 33: XIX-5). See Fig. 2A.
Utou, Otsukirai-mura, Kesen-gun ('^'(llj^j^S
51^ ^ B 3l ); Iwate, HONSHU; 39°07'N,
141°50'E; reported in questionnaire survey
conducted in 1 923 by K. Hasebe (Iwano, 1 974,
p. 56: XLV-4). See Fig. 2A.
Utougi, Ookawauchi-mura, Abe-gun (^"f^^PA
)Rlrt^W^7K); Shizuoka, HONSHU; 35°12'N,
138°22'E; reported in questionnaire survey
conducted in 1 923 by K. Hasebe (Iwano, 1 974,
p. 45:XXX-9). SeeFig. 2A.
Utsutsumi, Kamogawa-city (^IJIlTtTflM); Chiba,
HONSHU; 35°07'N, 140°04'E; collected 11
Nov. 1997-18 Dec. 1998 by PRIKU staff;
PRIKU, 19 (skeletons only). NE125.
Wachi-cho, Funai-gun (^p'^^^^fflJ); Kyoto,
HONSHU; ca. 35°17'N, 135°25'E; captured ca.
16 Apr. 1986, died 10 Jun. 1986; JMC, 1 (skull
184
FIELDIANA: ZOOLOGY
only). Blood samples collected before 1992 by
Nozawa et al. (1991, p. 414; 1996, p. 6).
SW4L
Wada-mura, Ooi-gun (AtS?P^B9^); Fukui,
HONSHU; 35°29TSf, 135°35'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 39: XXV-5). See Fig.
2A.
Wainai, Kariya-mura, Shimohei-gun (TPB'^fP
XlJM^W^I^); Iwate, HONSHU; 39°41'N,
141°43'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 56:XLV-10). SeeFig. 2A.
Wakahowatauchi, Nagano-city (:^i?rfT^#;^
rt); Nagano, HONSHU; 36°37'N, 138°16'E;
collected 17 Nov. 1998 by PRIKU staff;
PRIKU, 1 (skeleton only). NE50.
Wakamiya-Hachimanjinja, Takayama-mura,
Kamitakai-gun {hm^^m^\l\^^'U R^^
tt); Nagano, HONSHU; 36°41'N, 138°25'E
collected 5 Feb. 2000 by M. Aimi; PRIKU, 1
(skeleton only). NE49.
Wakasa-cho, Yazu-gun (AHiPS^fflT); Tottori,
HONSHU; ca. 35°20'N, 134°24'E; captive
group translocated from Wakasa to PRIKU,
mtDNA samples collected ca. 1986-1991 by
Hayasaka et al. (1991, p. 400). Blood samples
collected before 1992 by Nozawa et al. (1991,
p. 414; 1996, p. 6). Microsatellite DNA ana-
lyzed before 1996 by Domingo-Roura et al.
(1997, p. 358). External measurements taken
before 1997 by Hamada et al. (1996a, pp. 98,
99). MtDNA samples collected before 2003 by
Y. Kawamoto (2002, p. 60). SW50.
Wake vicinity, Kamikawa-mura, Minami-
muro-gun (^#ftlP± Jl| W^-^ffe); Mie,
HONSHU; 33°49'N, 135°55'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 38: XXIII-55). See
Fig. 2A.
Wakigahata-mura, Inukami-gun (:^-b^flS '^'"llffl
^); Shiga, HONSHU; 35°13'N, 136°22'E; re-
ported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 38:
XXIV-21). SeeFig. 2A.
Wakinosawa-mura vicinity, Shimokita-gun (Tdb
?P fiS i? )H W - w ); Aomori; HONSHU;
41°09'N, 140°46'E; collected 10 Oct. 1961 by
PRIKU staff; PRIKU, 1 (skull only). Reported
before 1970 by C. B. Koford (1969, p. 12; cf.
Azuma, 1985, p. 3; Fooden & Aimi, 2003, p.
111). Blood samples collected before 1992 by
Nozawa et al. (1991, p. 414; 1996, p. 6). NE5.
Warabitaira, Takayama-mura, Kamitakai-gun (Jb
i^#iPi^Uj;^M¥); Nagano, HONSHU;
36°41'N, 138°24'E; collected 3 Jan. 2000 by M.
Aimi; PRIKU, 1 (skeleton only). NE49.
Wasedasuirin vicinity. Shionomachi-mura, Iwa-
fune-gun {^li^^^m^^^mm^^^;
Niigata, HONSHU; 38°25'N, 139°34'E; re-
ported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 48:
XXXII-24). See Fig. 2A.
Wataka vicinity, Owase-cho, Kitamuro-gun (db
#«lPMiKfflI}lgEffe); Mie, HONSHU;
34°04'N, 136°I1'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 37: XXIII-46). See Fig. 2A.
Watarai-gun (Jg^lP); Mie, HONSHU; ca.
34°20'N, 136°30'E; acquired before 1939 by N.
Kuroda (1938, p. 112; 1940, pp. 270, 271);
formerly in private collection of N. Kuroda,
destroyed by fire in 1945 (Austin et al., 1948,
p. 4), 1 (not seen). Not mapped (see Oo-
miya-cho).
Yabitsuyama National Forest, Kanita-mura, Hi-
gashitsugaru-gun (^/$^IPMffl^^ffllll@
W^); Aomori, HONSHU; 41°03'N, 140°39'E;
reported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 57:
XLVI-16). SeeFig. 2A.
Yaheishiroburaku, Okugawa-mura, Yama-gun
(IP0?PIIJI|^?^^¥Eg^P^^); Fukushima,
HONSHU; 37°43'N, 139°41'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 52: XLI-6). See Fig.
2A.
Yaichirousawa vicinity, Oohata-mura, Shi-
mokita-gun (T:lblPA;lffl^?^^-fiP)Rflb);
Aomori, HONSHU; 41°24'N, 141°08'E; re-
ported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 57:
XLVI-19). SeeFig. 2A.
Yakeyama, Azusagawa-mura, Minamiazumi-gun
(lt^«?P^JI|^:^aj); Nagano, HONSHU;
36°11'N, 137°49'E; collected 7-10 Nov. 1998
by PRIKU staff; PRIKU, 5 (skeletons only,
with fragmented skulls). NE94.
FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES
185
Yakeyama vicinity, Umegashima-mura, Abe-gun
( ^ ^ ?P ^S <r 1 ;t4 'Jt^ 111 fife ); Shizuoka,
HONSHU; 35°16'N, 138°20'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 45: XXX-8). See Fig.
2A.
Yaku vicinity ( M ^ A ); Kagoshima,
YAKUSHIMA; ca. 30°16'N, 130°29'E; col-
lected ca. 1989 by S. Azuma; PRIKU, 4
(skeletons only). 5^/07.
Yaku-cho, Kumage-gun (^^?PM:^fflI); Ka-
goshima, YAKUSHIMA; coordinates un-
known; blood samples collected before 1992
by Nozawa et al. (1991, p. 414; 1996, p. 7).
Not mapped.
Yakushima ( M ^ S ); Kagoshima, YAKU-
SHIMA; 30°13'-30°27'N, 130°23'-130°40'E;
collected 4-18 Oct. 1904 by A. Owston;
BM(NH), 5. Collected 2 Jul. 1905 by A. Ow-
ston (Kuroda, 1940, p. 273; Imaizumi, 1962, p.
10); formerly in private collection of N. Ku-
roda (destroyed by fire in 1945), 1 (skin
only[?]). Captured alive, died in captivity 1 1
Oct. 1960; JMC, 1 (skeleton only). Field sur-
vey conducted 1973-1974 by 11 teams of in-
vestigators (Ashizawa, 1983, p. 10). Captives,
specimens obtained 29 Jan. 1986-4 Feb. 2000;
PRIKU, 32 (skeletons only, including 5 with
fragmented skulls and 4 with postcranials
only). MtDNA samples collected before 1987
by Hayasaka et al. (1986, p. 346). Microsatel-
lite DNA analyzed before 1996 by Do-
mingo-Roura et al. (1997, p. 358). External
measurements taken before 1997 by Hamada
et al. (1996a, pp. 98 ,99). Not mapped.
Yakushima, Census Area 1 (Nagata-Issou),
Kamiyaku-cho, Kumage-gun C^^W-^M^
mj^Ba- -M); Kagoshima, YAKUSHIMA;
30°22.5'N, 130°24'E-30°27'N, 130°30'E; 25
troops censused 23-25 Jul. 1991 by Yoshihiro
et al. ( 1 998, p. 1 80). Not mapped.
Yakushima, Census Area 2 (Issou-Miyanoura),
Kamiyaku-cho, Kumage-gun (^^?P_hM^
Urr - ^^ - "g (7) 5i ); Kagoshima,
YAKUSHIMA; 30°27'N, 130°30'E-30°25'N,
130°34'E; 14 troops censused 27-30 Jul. 1991
by Yoshihiro et al. (1998, p. 180). Not
mapped.
Yakushima, Census Area 3 (Miyanoura-Ambo)
(^(D)i- '^my, Kagoshima, YAKUSHIMA;
30°24.5'N, 130°34.5'E-30°19'N, 130°39.5'E; 8
troops censused 27 Jul.-2 Aug. 1992 by Yo-
shihiro et al. (1998, p. 180). Not mapped.
Yakushima, Census Area 4 (Ambo-Onoaida),
Yaku-cho, Kumage-gun (B^^BX^'^M
- M :^f^ ); Kagoshima, YAKUSHIMA;
30°19'N, 130°39.5'E-30°14.5'N, 130°33'E; 19
troops censused 2^ Aug. 1991 by Yoshihiro
et al. (1998, p. 180). Not mapped.
Yakushima, Census Area 5 (Onoaida-Kurio),
Yaku-cho, Kumage-gun (M^W^MX^M:^
m - ^ ^ ); Kagoshima, YAKUSHIMA;
30°14.5'N, 130°33'E-30°16'N, 130°26'E; 24
troops censused 16-23 Jul. 1992 by Yoshihiro
et al. (1998, p. 180). Not mapped.
Yakushima, Census Area 6 (Kurio-Segire) (H^
- MW); Kagoshima, YAKUSHIMA; 30°16'N,
130°26'E-30°19'N, 130°24.5'E; 17 troops
censused 29 Jul.-13 Aug. 1993 by Yoshihiro
et al. (1998, p. 180). Not mapped.
Yakushima, Census Area 7 (Hanyama-Segire),
Kamiyaku-cho, Kumage-gun (^^^-hM^
rr¥Uj- MWy, Kagoshima, YAKUSHIMA;
30°19'N, 130°24.5'E-30°22.5'N, 130°24'E; 24
troops censused 2-7 Aug. 1990 by Yoshihiro
et al. (1998, p. 180; cf Yoshihiro et al., 1999,
p. 41 1). Not mapped.
Yakushima, Loc. A ( M ^ A ); Kagoshima,
YAKUSHIMA; ca. 30°24'N, 130°24'E;
mtDNA samples collected before 2003 by S.
Hayaishi and Y. Kawamoto (2002, p. 164).
SWIOI.
Yakushima, Loc. B ( M ^ S ); Kagoshima,
YAKUSHIMA; ca. 30°23'N, 130°23'E;
mtDNA samples collected before 2003 by S.
Hayaishi and Y. Kawamoto (2002, p. 164).
SWlOl.
Yakushima, Loc. C ( M ^ S ); Kagoshima,
YAKUSHIMA; ca. 30°23'N, 130°23'E;
mtDNA samples collected before 2003 by S.
Hayaishi and Y. Kawamoto (2002, p. 164).
SWJOl.
Yakushima, Loc. D ( M ^ A ); Kagoshima,
YAKUSHIMA; ca. 30°22'N, 130°23'E;
mtDNA samples collected before 2003 by S.
Hayaishi and Y. Kawamoto (2002, p. 164).
SWIOI.
Yakushima, Loc. E ( M ^ ft ); Kagoshima,
186
FIELDIANA: ZOOLOGY
YAKUSHIMA;
mtDNA samples
Hayaishi and Y.
SWIOI.
Yakushima, Loc.
YAKUSHIMA;
mtDNA samples
Hayaishi and Y.
SWlOl.
Yakushima, Loc.
YAKUSHIMA;
mtDNA samples
Hayaishi and Y.
SWIOI.
Yakushima, Loc.
YAKUSHIMA;
mtDNA samples
Hayaishi and Y.
SWIOI.
Yakushima, Loc.
YAKUSHIMA;
mtDNA samples
Hayaishi and Y.
SWIOI.
Yakushima, Loc.
YAKUSHIMA;
mtDNA samples
Hayaishi and Y.
SWIOI.
Yakushima, Loc.
YAKUSHIMA;
mtDNA samples
Hayaishi and Y.
SWIOI.
Yakushima, Loc.
YAKUSHIMA;
mtDNA samples
Hayaishi and Y.
SWIOI.
Yakushima, Loc.
YAKUSHIMA;
mtDNA samples
Hayaishi and Y.
SWIOI.
Yakushima, Loc.
YAKUSHIMA;
mtDNA samples
Hayaishi and Y.
SWIOI.
ca. 30°22'N, 130°23'E;
collected before 2003 by S.
Kawamoto (2002, p. 164).
F ( M ^ fi ); Kagoshima,
ca. 30°21'N, 130°24'E;
collected before 2003 by S.
Kawamoto (2002, p. 164).
G { M ^ M )', Kagoshima,
ca. 30°19'N, 130°24'E;
collected before 2003 by S.
Kawamoto (2002, p. 164).
H ( M ^ ft ); Kagoshima,
ca. 30°18'N, 130°26'E;
collected before 2003 by S.
Kawamoto (2002, p. 164).
I ( M ^ fi ); Kagoshima,
ca. 30°23'N, 130°24'E;
collected before 2003 by S.
Kawamoto (2002, p. 164).
J ( M ^ fi ); Kagoshima,
ca. 30°23'N, 130°24'E;
collected before 2003 by S.
Kawamoto (2002, p. 164).
K ( M ^ S ); Kagoshima,
ca. 30°23'N, 130°24'E;
collected before 2003 by S.
Kawamoto (2002, p. 164).
L ( M ^ fi ); Kagoshima,
ca. 30°20'N, 130°25'E;
collected before 2003 by S.
Kawamoto (2002, p. 164).
M ( M ^ fi ); Kagoshima,
ca. 30°20'N, 130°26'E;
collected before 2003 by S.
Kawamoto (2002, p. 164).
N ( M y^ fi ); Kagoshima,
ca. 30°20'N, 130°27'E;
collected before 2003 by S.
Kawamoto (2002, p. 164).
Yakushima, Loc. O {M X M ); Kagoshima,
YAKUSHIMA; ca. 30° 19^ 130°25'E;
mtDNA samples collected before 2003 by S.
Hayaishi and Y. Kawamoto (2002, p. 164).
SWIOI.
Yakushima, Loc. P ( M ^ ^ ); Kagoshima,
YAKUSHIMA; ca. 30°26'N, 130°32'E;
mtDNA samples collected before 2003 by S.
Hayaishi and Y. Kawamoto (2002, p. 164).
SWIOO.
Yakushima, Loc. Q ( M ^ S ); Kagoshima,
YAKUSHIMA; ca. 30°23'N, 130°34'E;
mtDNA samples collected before 2003 by S.
Hayaishi and Y. Kawamoto (2002, p. 164).
SWIOO.
Yakushima, Loc. R ( M ^ S ); Kagoshima,
YAKUSHIMA; ca. 30°19'N, 130°36'E;
mtDNA samples collected before 2003 by S.
Hayaishi and Y. Kawamoto (2002, p. 164).
SW102.
Yakushima, Loc. S ( M ^ ^ ); Kagoshima,
YAKUSHIMA; ca. 30°18'N, 130°35'E;
mtDNA samples collected before 2003 by S.
Hayaishi and Y. Kawamoto (2002, p. 164).
SW102.
Yakushima, Loc. T ( M. X M ); Kagoshima,
YAKUSHIMA; ca. 30°16'N, 130°35'E;
mtDNA samples collected before 2003 by S.
Hayaishi and Y. Kawamoto (2002, p. 164).
SWI02.
Yakushima, Loc. U ( M ^ B ); Kagoshima,
YAKUSHIMA; ca. 30°15'N, 130°30'E;
mtDNA samples collected before 2003 by S.
Hayaishi and Y. Kawamoto (2002, p. 164).
SWI02.
Yakushima, southwestern (M^B^SnP); Ka-
goshima, YAKUSHIMA; ca. 30° 19^
130°28'E; mtDNA samples collected before
2003 by Y. Kawamoto (2002, p. 60). SWIOI.
Yakushima, western, road vicinity, high zone (M
XBMn^WMmt^w), 800-1199 m elevation;
Kagoshima, YAKUSHIMA; ca. 30°20'N,
130°25.5'E; fecal samples collected for dietary
analysis, 1999-2001, by G. Hanya et al.
(2003a, p. 52). SWIOI.
Yakushima, western, road vicinity, low zone (M
XB'^^Pl^M.i^t^^), 0-399 m elevation;
Kagoshima, YAKUSHIMA; ca. 30°18'N,
130°25'E; fecal samples collected for dietary
FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES
187
analysis, 1999-2001, by G. Hanya et al.
(2003a, p. 52). SWIOl.
Yakushima, western, road vicinity, middle zone
(MXM'^^WM^f^w), 400-799 m eleva-
tion; Kagoshima, YAKUSHIMA; ca. 30°19'N,
130°25.5'E; fecal samples collected for dietary
analysis, 1999-2001, by G. Hanya et al.
(2003a, p. 52). 5^^707.
Yakushimayama, Kamiyaku-mura, Kumage-gun
{M^W±MX^MXB[hy, Kagoshima,
YAKUSHIMA; 30°24'N, 130°31'E; reported
in questionnaire survey conducted in 1923 by
K. Hasebe (Iwano, 1974, p. 12: I-l). See Fig.
2A.
Yamabuki, Takamori-machi, Shimoina-gun (~F
^M^m^^iU^y, Nagano, HONSHU;
35°34'N, 137°52'E; collected 2 Dec. 1998 by
PRIKU staff; PRIKU, 1 (skeleton only). NE76.
Yamada, Yamana-mura, Hata-gun (tH^lPLil^
^ Oj ffl ); Kochi, SHIKOKU; 33°01'N,
132°48'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 21: X- 10). See Fig. 2 A.
Yamada-mura, Kamitakai-gun ( _L S ^ IP ill B9
^y, Nagano, HONSHU; 36°42'N, 138°27'E;
reported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 46:
XXXI-28). See Fig. 2A.
Yamadera, Yamagata-city (|JL| Jf^TfT |Jj ^); Ya-
magata, HONSHU; ca. 38°18'N, 140°26'E;
mtDNA samples collected before 1998 by Y.
Kawamoto (1997, p. 33; 2002, p. 60). NE28.
Yamagata-mura, Kunohe-gun (APlPlljff^^);
Iwate, HONSHU; 40°08'N, 14r35'E; mtDNA
extracted from hunting trophies in Dec. 1998
by T. Agatsuma and M. Ishigami (2002, p. 81).
NE12.
Yamanaka, Futtsu-city ( S /$ TfT |JL| 4" ); Chiba,
HONSHU; 35°09'N, 139°55'E; collected 30
Mar. 1999 by PRIKU staff; PRIKU, 2 (skele-
tons only). NE120.
Yamate vicinity, Youra-mura, Kuma-gun (^M
lPEg}iWUj¥flfe); Kumamoto, KYUSHU;
32°19'N, 130°52'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 16: IV-5). See Fig. 2A.
Yamavvakiyama, Sogo-mura, Nakatado-gun {W
^J^iP+iPWOjiESaj); Kagawa, SHIKOKU;
34°08'N, 133°52'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 24: XII- 1). See Fig. 2 A.
Yanagisawa, Nakagawa-mura, Kamiina-gun (Ji
^M^^}\\^^yRy Nagano, HONSHU;
35°37'N, 137°57'E; collected 7 Jul. 1998 by M.
Aimi; PRIKU, 1 (skeleton only). NE71.
Yanagitani, Kaiinji-mura, Otokuni-gun (ZlllIlP
M^li^^^^y Kyoto, HONSHU; 34°55'N,
135°40'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 36: XXII-27). See Fig. 2A.
Yanagitani, Takahoko-son, Katsuura-gun (05^
IPSi^W^^); Tokushima, SHIKOKU;
33°56'N, 134°25'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 23: XI-16). See Fig. 2A.
Yarigatake { % T ^ \ 3180 m; Nagano;
HONSHU; 36°21'N, 137°39'E; observed on
summit before 1995 by S. Izumiyama et al.
(2003, p. 466; pers. comm., 29 Dec. 2003).
NE88.
Yashiroyama, Niiya, Yashiro-mura, Nima-gun
( M S ?P A ft W if M A <t OJ ); Shimane,
HONSHU; 35°03'N, 132°27'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 28: XVI-28). See Fig.
2A.
Yasui National Forest, Tomioka-mura,
Agawa-gun (a^JII?PS[^W$MSWW);
Kochi, SHIKOKU; 33°40'N, 133°15'E; re-
ported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 22: X-23).
See Fig. 2A.
Yatakiyama vicinity, Nishida, Yusato-mura,
Nima-gun (SS^PMM^® BQ ^)i Oj ffe);
Shimane, HONSHU; 35°06'N, 132°24'E; re-
ported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 28:
XVI-29). See Fig. 2A.
Yatsudake vicinity, Tomiyama-mura, Kitashi-
tara-gun (dblS^^PsUj^A^i^ffe); Aichi,
HONSHU; 35°10'N, 137°48'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 40: XXVII-7). See
Fig. 2A.
Yatsugatake National Forest, Kitamaki-mura,
Minamisaku-gun (^^XWit'^'^ fX'r ^^
W#); Nagano, HONSHU; 36°03'N, 138°28'E;
reported in questionnaire survey conducted in
188
FIELDIANA: ZOOLOGY
1923 by K. Hasebe (Iwano, 1974, p. 46:
XXXI-36). See Fig. 2A.
Yatsugatake National Forest, Minamimaki-mura,
Minamisaku-gun {'^i^X^'Mi^X^A'r ^
1 W ^ ); Nagano, HONSHU; 35°59'N,
138°27'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 46: XXXl-38). See Fig. 2A.
Yedo i\Lp). See Tokyo.
Yo-mura, Higashimuro-gun ( !^ # ft ^ E3 ^ );
Wakayama, HONSHU; 33°49'N, 135°45'E;
reported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 34:
XX-49). See Fig. 2A.
Yogo-cho, Ika-gun (^#^^^^1); Shiga,
HONSHU; ca. 35°32'N, 136°13'E; mtDNA
samples collected before 2003 by Y. Kawa-
moto (2002, p. 60). SW5.
Yokogawaburaku vicinity, Togouchi-son, Ya-
magata-gun (UimiPFMrt^WJH^P^^jfi);
Hiroshima, HONSHU; 34°40'N, 132°20'E;
reported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 26:
XIV-5). See Fig. 2A.
Yokohama-city (IH^T^) (= Jokohama); Kana-
gawa, HONSHU; 35°27'N, 139°39'E; 118
skulls purchased 1900-1902 by K. A. Haberer,
collection locality unknown (cf. Schweyer,
1909, p. 8); AIUM, 32 (skulls only); ZSBS, 1
(skull only). Not mapped.
Yokokabe National Forest vicinity, Nagano-
hara-mura, Agatsuma-gun (n"$^Mi?J^^t
^MBW^fte); Gunma, HONSHU; 36°32'N,
138°39'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 51:XL-9). SeeFig.2A.
Yokomichi vicinity, Nichihara-mura, Kano-
ashi-gun (E£^BJ^4^^31'te); Shimane,
HONSHU; 34°32'N, 131°49'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 28: XVI-9). See Fig.
2A.
Yokootsuzukiyama vicinity, Hane-mura,
Aki-gun (^SiP^^Wmmaajffe); Kochi,
SHIKOKU; 33°22'N, 134°06'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 22: X-46). See Fig.
2A.
Yokose, Ootaki-machi, Isumi-gun (^P^?PA^
SKT1t)i); Chiba, HONSHU; 35°irN,
140°10'E; collected 22 Jun.-ll Dec. 1997 by
PRIKU staff; PRIKU, 3 (skeletons only, in-
cluding 1 with fragmented mandible). NE126.
Yokoze-machi, Chichibu-gun (^^IPIt^fflJ);
Saitama, HONSHU; ca. 35°58'N, 139°07'E;
mtDNA samples collected before 1998 by Y.
Kawamoto (1997, p. 33; 2002, p. 60). NEJ17.
Yonezawa, Tomiyama-machi, Awa-gun ($M1P
ailjffll7K)H); Chiba, HONSHU; 35°04'N,
139°54'E; collected 30 Mar. 1999 by PRIKU
staff; PRIKU, 1 (skeleton only). NEJ21.
Yoshida-mura, lishi-gun (ISS^PnBa^); Shi-
mane, HONSHU; 35°11'N, 132°55'E; reported
(as possibly temporary habitat) in question-
naire survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 28: XVI-34). Collected 7 Jul.
1995 by PRIKU staff; PRIKU, 1 (skeleton
only). SW59.
Yoshigo shell mound, Tahara-cho, Atsumi-gun
()lM?PfflJ^fflI^^Mii); Aichi, HONSHU;
coordinates unknown; Late Jomon subfossil
(age >2.3 Ka) reported before 2003 by M. Ku-
roda (2002a, p. 115); Kyoto University An-
thropology Laboratory, 1 (mandible only; not
seen). Not mapped.
Yotsugadake vicinity, Aiuchi-mura, Ki-
tatsugaru-gun (db5*g?P*l rt^EB 7 Sftfe);
Aomori, HONSHU; 41°03'N, 140°22'E; re-
ported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 57:
XLVI-11). SeeFig.2A.
Youkurayama, Funaki-son, Toyota-gun (SBB^
^r' ^ ^ ffl ^ UJ ); Hiroshima, HONSHU;
34°28'N, 132°59'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 26: XIV-23). See Fig. 2A.
Youkurayama, Kamikitakata-son, Toyota-gun
(^m^±ity5^mt\h); Hiroshima,
HONSHU; 34°26'N, 132°58'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 26: XIV-22). See Fig.
2A.
Youkurayama, Ookawa-son, Toyota-gun (WBB
^AM^MtiU); Hiroshima, HONSHU; co-
ordinates unknown; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 26: XIV-24). Not mapped.
Youkurayama, Zen-nyuji-son, Toyota-gun (MB9
FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES
189
?P#A#^ffl^UL|); Hiroshima, HONSHU;
34°26'N, 132°56'E; reported in questionnaire
sur\'ey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 26: XIV-21). See Fig. 2A.
Youzenotaki, Fuka-son, Kawakami-gun (JI|_hlP
S 1^ ^ ffl ^ y >i ); Okayama^ HONSHU:
34°46'N. 133°29'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 27: XV- 1). See Fig. 2 A.
Yudomari, Yaku-cho, Kumage-gun (^^IPM^
BI ^ )S ); Kagoshima, " YAKUSHIMA:
30°15'N,130°29'E; collected 25 Feb. 1989 by
PRIKU staff; PRIKU, 18 (skeletons only, in-
cluding 1 postcranials only). SW102.
Yugawara-machi, Ashigarashimo-gun (S:ffiT^
^ )RI J^ fflj ); Kanagmva, HONSHU; ca.
35°09'N, 139°04'E; provisioning initiated in
1956 (Study Group on the Present Status of
Japanese Monkeys, 1977b, p. 23). Blood sam-
ples collected before 1992 by Nozawa et al.
(1991, p. 414; 1996, p. 6). MtDNA samples
collected before 1998 by Y. Kawamoto (1997,
p. 33). NEWS.
Yukihikoyama, Kaya-mura, Shikama-gun {^M
iPE^^l^Ul); Hyogo, HONSHU;
34°58'N, 134°4rE; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 30: XVIII- 13). See Fig. 2 A.
Yumigiyama, Shiiba-son, Nishiusuki-gun (®Q
tfiP^^^^TKlij); Miyazaki, KYUSHU;
32°29'N, nriO'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 15: 11-32). See Fig. 2 A.
Yumoto, Yubara-cho, Maniwa-gun (KfilPi^J^
fflJ^^^); Okayama, HONSHU; 35°13'N,
133°45'E; reported in questionnaire sur\ey
conducted in 1923 by K. Hasebe (Iwano, 1974.
p. 27:XV-14). SeeFig. 2A.
Yunodake, Oohira, Ryogauchi-mura, Ihara-gun
(^iilPia^)Rlrt^A¥^i?^); Shizuoka,
HONSHU; 35°09'N, 138°27'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano, 1974, p. 45: XXX- 10). See Fig.
2A.
Yunoyama, Komono-cho, Mie-gun {E-M-WM.^
fflj ^^ y OJ ); Mie, HONSHU; 35°01'N,
136°27'E; reported present before 1939 by N.
Kuroda(1938, p. 112; 1940, p. 270). SPF/^.
Yunoyama, Komono-mura, Mie-gun (Hj
i? ^ ^^ y OJ ); Me, HONSHU; 35°01'N,
136°27'E; reported in questionnaire survey
conducted in 1923 by K. Hasebe (Iwano, 1974,
p. 36: XXIII-5). See Fig. 2A.
Yuumayama, Kawata-son, Oe-gun (^^^JHffl
^ i^ ^ U \h ): Tokushima, SHIKOKU;
34°02'N, 134°14'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 23: XI-26). See Fig. 2 A.
Yuzuruhayama, Nada-mura, Mihara-gun (HJ^
iPH^MS^LiJ); Hyogo, AWA^JISHIMA;
34°14'N, 134°49'E; reported in questionnaire
survey conducted in 1923 by K. Hasebe
(Iwano, 1974, p. 30: XVlII-1). See Fig. 2A.
Zarugadake vicinity, Suzurishima-mura, Mi-
namikoma-gun {'h^W-^WM'¥i^T ^^);
Yamanashi. HONSHU; 35°22'N, 138°19'E;
reported in questionnaire survey conducted in
1923 by K. Hasebe (Iwano, 1974, p. 48:
XXXIII-6). See Fig. 2A.
Zoukiyama, Higashikawa-mura, Higashikan-
bara-gun (^^J^IP^JH^MtKUj); Niigata,
HONSHU; 37°35'N. 139°26'E; reported in
questionnaire survey conducted in 1923 by K.
Hasebe (Iwano. 1974, p. 47: XXXII- 17). See
Fig. 2A.
190
FIELD! AN A: ZOOLOGY
Appendix 5. Local Variation in Sitting Height (mm) in Macaca fuscata Adults on Honshu (cf. Fig. 5)'
Lati-
tude
Longitude (E)- '
(N)-
132°
133'
J
137° 138°
139°
140°
Adult females
38°
550.0
37°
550-550 (2)
36°
573.0 ± 42.00 557.6 ± 34.81
35°
551.3 ± 50.33
525-603 (J) 510-608(7)
564.5 ± 22.61 558.7 ± 20.65
521.0
528.0 ± 27.48
34°
515-625(4)
547.5 ± 29.68
497-590(79)-'
542.0 ±
18.81
523-604 (i7) 535-573 (J)
Adult males
512-530(2)
487-545 (4)
38°
594
37°
(/)
36°
603.6 ± 32.53 612.5 ± 29.02
35°
615
552-630 (5) 553-648 (S)
623.2 ± 25.66 623
565
581.0 ± 17.67
34°
(/)
577.8 ± 40.23
504-625 (ISf
584.9 ±
(7)'
39.91
570-682 {22) (/)
U)
551-614(70)
' Data derived from specimens examined, except as otherwise indicated in footnotes 5 and 6. Within each cell,
figures indicate mean ± SD (where n > 2). extremes, and sample size (italicized, in parentheses).
- Column and row headings indicate lower limits of one-degree latitude-longitude blocks.
^ No data are available for specimens originating in the longitudinal zone 134-136.99°E.
^ Iwamoto (1971, table opp. p. 156) has published the following data for two additional localities situated within
this latitude-longitude block: Kochi (provisioned group), 530.0 ± 28.04 mm, n = 72; Hasumi (nonprovisioned group),
541.4 ± 13.39 mm, n = 5.
'' Data from Iwamoto (1971, table opp. p. 156); extreme values not published.
'' Includes one adult male (sitting height = 595 mm) measured at Kochi (38°28'N, 132°53'E; provisioned group)
by Iwamoto (1971, table opp. p. 156).
FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES 191
Appendix 6. Local Variation in Anterior Trunk Length (mm) in Macaca fuscata Adults on Honshu (cf. Fig. 6)'
Lati- Longitude (E)-
tude
(N)- 132° 133° 134° 135°
Adult females
36°
35° 381.2 ± 23.19
363-420 (5)
361.4 ± 12.26(7) 358.5 ± 16.69(27)^ 371.8 ± 19.53(47)-^
34° 373.4 ± 26.13
332-415 {14)
364.0 ± 11.24(6)^
372.0 ± 21.15(46)-'
Adult males
36°
35° 385
(/)
397.6 ± 12.01 {Sy 411.2 ± 23.88(77)-^
34° 364.4 ± 37.25
315-435(72)
394.5 ± 20.12(6)^
399.1 ± 16.68(22)'
' Within each cell, figures above the dashed line indicate data derived from specimens examined — mean ± SD
(where n > 2), extremes, and sample size (italicized, in parentheses): figures below the dashed line indicate data
published by Hamada et al. (1996a, pp. 101. 102) — mean ± SD and sample size. To conserve space, data for three
localities on Honshu north of 37°N are not included in the body of this table; these data are as follows: Takane,
38°20'N, 139°37'E, female, anterior trunk length = 371 mm, n = 1; Komogawa, 38°18'N, 139°36'E, female, anterior
trunk length = 371 mm, n = 1, male, anterior trunk length = 399 mm, n = 1: Shimokita, 41°08'N, 140°50'E, females,
367.9 ± 22.47 mm. n = 10 (Hamada et al., 1996a. p. 102).
- Column and row headings indicate lower limits of one-degree latitude-longitude blocks.
"* Captive and/or provisioned.
•» Mihara-city, 34°24'N, 133°05'E.
■'' Gagyusan. 34°49'N, 133°37'E (translocated to Nagatoro).
* References: Aizawa and Hagiwara (2001, pp. 5-6); M. Hagiwara (pers. comm., 3 Oct. 2002); these measurements
were originally reported as head and body length.
192 FIELDIANA: ZOOLOGY
Appendix 6. Extended.
Longitude (E)^
136° 137° 138° 139° 140°
Adult females
396.0 ± 53.14 369.6 ± 26.94 374.4 ± 11.19
344-470(4) 338-414(7) 355-392(77)
394.4 ± 13.42(77) 391.3 ± 20.30(42) 371.4 ± 12.07(76)
384.1 ± 20.34 380.7 ±23.16 347.5 358.6 ± 13.26
329-428 (J7) 355-400 (J) 335-360(2) 329-371(9)
369.9 ± 21.24(75)3 346.9 ± 13.82 (72)^
374.6 ± 17.76(57)
Adult males
400.4 ± 25.53
415.4 ± 21.66
406.0 ± 4.36
371-434(5)
383-446 (9)
403-411 (3)
413.6 ± 20.44(29)
403.3 ± 18.42(70)
411.5 ± 18.17
401
380
394.7 ± 23.23
375-441 (22)
(1)
(1)
365-450 (13)
404.2 ± 13.44(74)3 402.0(2)"
401.1 ± 19 A9 (34)
FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES 193
DXl
^
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FIELDIANA: ZOOLOGY
a
a
pecimens
alities on
T/ATL =
4.15 mm
0
ON
'^ 0
f^
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g u II '^.
c; <u —
73^=^
"■^ '^ <-" II
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"S'S E^
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Hamada et al.
e data are as fo
n = 1, T/ATL
tude-longitude 1
te6).
•S ^i E 1 S
-lio i 'S
.- t/5 .C On 60
u L3 ca <u . r^
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e data
dy of 1
il leng
102. 1
ts of c
i Naga
(see T
le.
w*
E •- ,, ,r n.— -a ^Tz:
^-ooju"", <o ^ a
"^
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: mean ± SD (where n >
ures below the dashed li
of 37°N are not included
gawa, 38°18'N, 139°36'E
19.4% (Hamada et al.. 1
id row headings indicate
d/or provisioned.
y. 34°24'N, 133°05'E.
34°49'N. 133°37'E(trans
Aizawa and Hagiwara. 2
n, anterior trunk length n
3.0 ± 7.95
1.0%
9.2 ± 10.4
4.8%
00 (N Ov (N
0^
+1^
t^^E" --Si 8^
't 't
' Tail len;
examined;
Honshu no
22.1%; Ko
(5), T/ATL
~ Column
3 Captive
'' Mihara-
' Gagyus,
^ Referen
^ Watarai
00 (N
01 n
FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES
195
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FIELDIANA: ZOOLOGY
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197
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198
FIELDIANA: ZOOLOGY
Index
abbreviations, institutional names 1
albinism 8
Allen's rule 85
antibodies to cedar pollen
antigens 43
arboreality 51
autosexual behavior 62
Bergmann's rule 11, 13, 14, 85
birth season 54
birth weight 65
birthrate 66
Blakiston's Line 83
blood proteins 37, 75, 85
collections studied 1
consortship 59
copulatory behavior 60
cranial measurements 17, 75, 85
geographic variation 19, 85
ontogeny 17, 19
provisioning, effect of 17
sexual dimorphism 17
dead infants, carried by mothers 68
dental emergence 19
dermatoglyphics 15
diagnosis
Macaco fuscata fuscata 8 1
Macaca fuscata yakui 83
diet 52
distribution, elevational 3, 50
distribution, geographic 3
Macaca fuscata fuscata 8 1
Macaca fuscata yakui 83
dominance rank and reproductive
success 60, 68
estrus 58
evolution and dispersal 83
extermination as agricultural
pest 5, 54
external measurements 8
anterior trunk length 10, 75, 85
body weight 13, 85
geographic variation 9
head and body length 8
hind foot length 12, 85
relative tail length 1 1
sexual dimorphism 9
sitting height 9, 75, 85
tail length 11, 85
fossils
macaque 72, 83
proboscidean 72, 83
genetic drift 39
geophagy 53
gestation period 63
glacial advances 83
Gloger's rule 8, 85
grooming 49
group
density 52
extinction 51
fission 51
size and composition 51
habitats 50, 85
hemoglobin amino-acid
sequence 42
history of scientific knowledge of
species 1
holotype, Macaca fuscata yakui 82
home range area 52
homosexual behavior 62
hybridization 61
implantation bleeding 64
inbreeding 54, 61
infant mortality 68
insular populations
extant 3
extinct 3
introduced 3
interbirth interval 66
junior synonym, first published
as 80
karyology 45
key to external characters 77
Korea Strait 83
land bridges 72, 83
lateral facial crest 5
lectotype designation 80
lifetime births per female 66
limb malformations 15
longevity 71
malaria 46
male emigration 54, 57
masturbation 62
mating season 54
menarche 57
menstrual cycle 58
milk secretion 69
mitochondrial DNA 22, 33, 35, 39,
75, 85
molting 5
multi-mount ejaculation 60
neonatal sex ratio 65
nomenclatural history, Macaca
fuscata 81
nonreproductive sexual behavior 62
nuclear DNA
intron 33
microsatellite 33
minisatellite 33
random amplified polymor-
phism 35
Y-chromosome 35
nursing 68
orbital measurements 20, 75
Palaeoloxodon naumanni 72
parasites 46
cestodes 47
lice 49
mites 49
nematodes 47
protistans 46
amebas 46
ci Hates 46
flagellates 46
sporozoans 46
Q fever bacterium 49
ticks 49
trematodes 47
viruses 49
paratypes, Macaca fuscata yakui 82
parturition 64
pelage 5
adult coloration 5. 75
geographic variation 6
hair banding 5
hair density 8
hair length 5
infantile coloration 5, 8
seasonal variation 5
population estimate 5
population growth rate 72
predators 54
reproductive senescence 71
rostral projection 21
sagittal crest 21
sea-level depressions 83
sexual skin 57
sexual maturation 57
side-whiskers. See lateral facial
crest
sika deer, association with 54
sleeping sites 51
Stegodon orientalis 72
subfossils 21
subspecies accounts
Macaca fuscata fuscata 11
Macaca fuscata yakui 82
subspecific recognition 72
suppressed name 78
surveys, geographic 3
swimming ability 51
synonymy
Macaca fuscata fuscata 77
Macaca fuscata yakui 82
syntypes, Macaca fuscata
fuscata 79
Tatar Strait 83
terrestriality 51
testis, maturation 57. 72
thermoregulation 8
Tsugaru Strait 83
twinning 65
type locality
Macaca fuscata fuscata 80
Macaca fuscata yakui 83
type series
Macaca fuscata fuscata 79
Macaca fuscata yakui 82
vernacular names 77
weaning 68
Yakushima-Sakhalin archipelago 83
INDEX
199
Note Added in Proof
Listed below are three rele\ ant publications that appeared while this monograph was in press.
FuJiTA. S.. H. Slgiira. F Mitslnaga. and K. Shinjizl. 2004. Hormone profiles and reproductive characteristics in
wild female Japanese macaques (Macaca fiiscata). .American Journal of Primatology. 64: 367-375.
R.\E, T. C. and T. Koppe. 2004. Holes in the head: Evolutionar> interpretations of the paranasal sinuses in catarrhines.
Evolutionary .Anthropology. 13: 211-223.
V'asev. p. L. 2004. Sex differences in sexual partner acquisition, retention, and harassment during female homosexual
consortships in Japanese macaques. American Journal of Primatology. 64: 397 — \09.
200 FIELDIANA: ZOOLOGY
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